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Gibbons () are apes in the family Hylobatidae (). The family historically contained one genus, but now is split into four extant genera and 20 species. Gibbons live in subtropical and tropical rainforest from eastern Bangladesh to Northeast India to southern China and Indonesia (including the islands of
Sumatra Sumatra is one of the Sunda Islands of western Indonesia. It is the largest island that is fully within Indonesian territory, as well as the sixth-largest island in the world at 473,481 km2 (182,812 mi.2), not including adjacent i ...
, Borneo and Java). Also called the lesser apes, gibbons differ from great apes (
bonobo The bonobo (; ''Pan paniscus''), also historically called the pygmy chimpanzee and less often the dwarf chimpanzee or gracile chimpanzee, is an endangered great ape and one of the two species making up the genus '' Pan,'' the other being the comm ...
s,
chimpanzee The chimpanzee (''Pan troglodytes''), also known as simply the chimp, is a species of great ape native to the forest and savannah of tropical Africa. It has four confirmed subspecies and a fifth proposed subspecies. When its close relative th ...
s, gorillas, orangutans and humans) in being smaller, exhibiting low sexual dimorphism, and not making nests. Like all apes, gibbons are tailless. Unlike most of the great apes, gibbons frequently form long-term pair bonds. Their primary mode of locomotion,
brachiation Brachiation (from "brachium", Latin for "arm"), or arm swinging, is a form of arboreal locomotion in which primates swing from tree limb to tree limb using only their arms. During brachiation, the body is alternately supported under each forelimb ...
, involves swinging from branch to branch for distances up to , at speeds as fast as . They can also make leaps up to , and walk bipedally with their arms raised for balance. They are the fastest of all tree-dwelling, nonflying mammals. Depending on the species and sex, gibbons' fur coloration varies from dark- to light-brown shades, and any shade between black and white, though a completely "white" gibbon is rare.


Etymology

The English word "gibbon" is a reborrowing from French and may originally derive from an
Orang Asli Orang Asli (''lit''. "first people", "native people", "original people", "aborigines people" or "aboriginal people" in Malay) are a heterogeneous indigenous population forming a national minority in Malaysia. They are the oldest inhabitants of ...
word.


Evolutionary history

Whole genome molecular dating analyses indicate that the gibbon lineage diverged from that of great apes around 16.8 million years ago (Mya) (95% confidence interval: 15.9–17.6 Mya; given a divergence of 29 Mya from Old World monkeys). Adaptive divergence associated with chromosomal rearrangements led to rapid radiation of the four genera 5–7 Mya. Each genus comprises a distinct, well-delineated lineage, but the sequence and timing of divergences among these genera has been hard to resolve, even with whole genome data, due to radiative speciations and extensive incomplete lineage sorting. An analysis based on morphology suggests that the four genera are ordered as (''Symphalangus'', (''Nomascus'', (''Hoolock'', ''Hylobates''))). A coalescent-based species tree analysis of genome-scale datasets suggests a phylogeny for the four genera ordered as (''Hylobates'', (''Nomascus'', (''Hoolock'', ''Symphalangus''))). At the species level, estimates from mitochondrial DNA genome analyses suggest that ''Hylobates pileatus'' diverged from ''H. lar'' and ''H. agilis'' around 3.9 Mya, and ''H. lar'' and ''H. agilis'' separated around 3.3 Mya. Whole genome analysis suggests divergence of ''H. pileatus'' from ''H. moloch'' 1.5–3.0 Mya. The extinct '' Bunopithecus sericus'' is a gibbon or gibbon-like ape, which until recently, was thought to be closely related to the hoolock gibbons.


Taxonomy

The family is divided into four
genera Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...
based on their
diploid Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectively ...
chromosome number: ''
Hylobates The genus ''Hylobates'' is one of the four genera of gibbons. Its name means "forest walker", from the Greek (, "forest") and (, "one who treads"). It was once considered the only genus, but recently its subgenera (''Hoolock'' ormerly ''Bun ...
'' (44), '' Hoolock'' (38), '' Nomascus'' (52), and '' Symphalangus'' (50). Also, three extinct genera currently are recognised: '' Bunopithecus'', '' Junzi'', and ''
Yuanmoupithecus ''Yuanmoupithecus'' is an extinct genus of gibbons that lived 8.2 to 7.1 million years ago during the late Miocene. It is currently the oldest gibbon known. It was discovered in Yuanmou, Yunnan Province, China China, officially the Peopl ...
''. Family Hylobatidae: gibbons * Genus '' Hoolock'' **
Western hoolock gibbon The western hoolock gibbon (''Hoolock hoolock'') is a primate from the gibbon family, Hylobatidae. The species is found in Assam, Mizoram, and Meghalaya in India, Bangladesh, and Myanmar west of the Chindwin River. Classification Mootnick and ...
, ''H. hoolock'' ** Eastern hoolock gibbon, ''H. leuconedys'' ** Skywalker hoolock gibbon, ''H. tianxing'' * Genus ''
Hylobates The genus ''Hylobates'' is one of the four genera of gibbons. Its name means "forest walker", from the Greek (, "forest") and (, "one who treads"). It was once considered the only genus, but recently its subgenera (''Hoolock'' ormerly ''Bun ...
'': dwarf gibbons **
Lar gibbon The lar gibbon (''Hylobates lar''), also known as the white-handed gibbon, is an endangered primate in the gibbon family, Hylobatidae. It is one of the better-known gibbons and is often kept in captivity. Taxonomy There are five subspecies of ...
or white-handed gibbon, ''H. lar'' ** Bornean white-bearded gibbon, ''H. albibarbis'' **
Agile gibbon The agile gibbon (''Hylobates agilis''), also known as the black-handed gibbon, is an Old World primate in the gibbon family. It is found in Indonesia on the island of Sumatra, Malaysia, and southern Thailand. The species is listed as endangere ...
or black-handed gibbon, ''H. agilis'' **
Western grey gibbon The western grey gibbon (''Hylobates abbotti''), also known as Abbot's grey gibbon, is a primate in the gibbon family, Hylobatidae. It was named after zoologist William Louis Abbott. Taxonomy Formerly, the western grey gibbon and northern grey ...
or Abbott's grey gibbon, ''H. abbotti'' ** Eastern grey gibbon or northern grey gibbon, ''H. funereus'' **
Müller's gibbon Müller's gibbon (''Hylobates muelleri''), also known as the southern grey gibbon, is a primate in the gibbon family, Hylobatidae. Taxonomy Formerly, the western grey gibbon (''H. abbotti'') and eastern grey gibbon (''H. funereus'') were consi ...
or southern grey gibbon, ''H. muelleri'' ** Silvery gibbon, ''H. moloch'' ** Pileated gibbon or capped gibbon, ''H. pileatus'' ** Kloss's gibbon, Mentawai gibbon or bilou, ''H. klossii'' * Genus '' Symphalangus'' ** Siamang, ''S. syndactylus'' * Genus '' Nomascus'': crested gibbons **
Northern buffed-cheeked gibbon The northern buffed-cheeked gibbon (''Nomascus annamensis'') is a newly discovered species of crested gibbon which is found in Vietnam, Cambodia, and Laos. Its habitat is in the humid subtropical and seasonal tropical forests of these countries. ...
, ''N. annamensis'' ** Concolor or black crested gibbon, ''N. concolor'' **
Eastern black crested gibbon The eastern black-crested gibbon (''Nomascus nasutus''), also known as the Cao-vit black crested gibbon or the Cao-vit crested gibbon, is a species of gibbon Gibbons () are apes in the family Hylobatidae (). The family historically contained o ...
or Cao Vit black crested gibbon, ''N. nasutus'' ** Hainan black crested gibbon, ''N. hainanus'' ** Northern white-cheeked gibbon, ''N. leucogenys'' ** Southern white-cheeked gibbon, ''N. siki'' ** Yellow-cheeked gibbon, ''N. gabriellae''


Extinct genera

* Genus '' Bunopithecus'' ** '' Bunopithecus sericus'' * Genus '' Junzi'' ** ''
Junzi imperialis ''Junzi imperialis'' is an extinct species of gibbon that was found in an Ancient Chinese noblewoman's tomb. The type species, based on an incomplete skull, was named ''Junzi imperialis'' in 2018 by Samuel Turvey and colleagues. It is believed t ...
'' *Genus ''
Yuanmoupithecus ''Yuanmoupithecus'' is an extinct genus of gibbons that lived 8.2 to 7.1 million years ago during the late Miocene. It is currently the oldest gibbon known. It was discovered in Yuanmou, Yunnan Province, China China, officially the Peopl ...
'' ** ''
Yuanmoupithecus xiaoyuan ''Yuanmoupithecus'' is an extinct genus of gibbons that lived 8.2 to 7.1 million years ago during the late Miocene. It is currently the oldest gibbon known. It was discovered in Yuanmou, Yunnan Province, China. The type species is ''Y. xiaoyuan'' ...
''


Hybrids

Many gibbons are hard to identify based on fur coloration, so are identified either by song or genetics. These morphological ambiguities have led to hybrids in zoos. Zoos often receive gibbons of unknown origin, so they rely on morphological variation or labels that are impossible to verify to assign species and subspecies names, so separate species of gibbons commonly are misidentified and housed together. Interspecific hybrids, within a genus, are also suspected to occur in wild gibbons where their ranges overlap. No records exist, however, of fertile hybrids between different gibbon genera, either in the wild or in captivity.


Description

One unique aspect of a gibbon's anatomy is the wrist, which functions something like a ball-and-socket joint, allowing for biaxial movement. This greatly reduces the amount of energy needed in the upper arm and torso, while also reducing stress on the shoulder joint. Gibbons also have long hands and feet, with a deep cleft between the first and second digits of their hands. Their fur is usually black, gray, or brownish, often with white markings on hands, feet and face. Some species such as the siamang have an enlarged throat sac, which inflates and serves as a resonating chamber when the animals call. This structure can become quite large in some species, sometimes equaling the size of the animal's head. Their voices are much more powerful than that of any human singer, although they are at best half a human's height. Gibbon skulls and teeth resemble those of the great apes, and their noses are similar to those of all catarrhine primates. The dental formula is .Myers, P. 2000
Family Hylobatidae
Animal Diversity Web. Accessed April 05, 2011-04-05.
The siamang, which is the largest of the 18 species, is distinguished by having two fingers on each foot stuck together, hence the generic and species names ''Symphalangus'' and ''syndactylus''.


Behavior

Like all primates, gibbons are social animals. They are strongly territorial, and defend their boundaries with vigorous visual and vocal displays. The vocal element, which can often be heard for distances up to , consists of a duet between a mated pair, with their young sometimes joining in. In most species, males and some females sing solos to attract mates, as well as advertise their territories. The song can be used to identify not only which species of gibbon is singing, but also the area from which it comes. Gibbons often retain the same mate for life, although they do not always remain sexually monogamous. In addition to extra-pair copulations, pair-bonded gibbons occasionally "divorce". Gibbons are among nature's best brachiators. Their ball-and-socket wrist joints allow them unmatched speed and accuracy when swinging through trees. Nonetheless, their mode of transportation can lead to hazards when a branch breaks or a hand slips, and researchers estimate that the majority of gibbons suffer bone fractures one or more times during their lifetimes. They are the fastest of all tree-dwelling, nonflying mammals. Attenborough, David. ''
Life of Mammals ''The Life of Mammals'' is a nature documentary series written and presented by David Attenborough, first transmitted in the United Kingdom from 20 November 2002. A study of the evolution and habits of the various mammal species, it was the four ...
'', "Episode 8: Life in the Trees", BBC Warner, 2003.
On the ground, gibbons tend to walk bipedally, and their Achilles tendon morphology is more similar to that of humans than that of any other ape.


Diet

Gibbons' diets are about 60% fruit-based,Gibbon - Monkey Worlds
Retrieved Feb-12-2015
but they also consume twigs, leaves, insects, flowers, and occasionally birds' eggs.


Genetics

Gibbons were the first apes to diverge from the common ancestor of humans and apes about 16.8 Mya. With a genome that has a 96% similarity to humans, the gibbon has a role as a bridge between Old World monkeys, such as
macaque The macaques () constitute a genus (''Macaca'') of gregarious Old World monkeys of the subfamily Cercopithecinae. The 23 species of macaques inhabit ranges throughout Asia, North Africa, and (in one instance) Gibraltar. Macaques are principally ...
s, and the great apes. According to a study that mapped synteny (genes occurring on the same chromosome) disruptions in the gibbon and human genome, humans and great apes are part of the same superfamily ( Hominoidea) with gibbons. The
karyotype A karyotype is the general appearance of the complete set of metaphase chromosomes in the cells of a species or in an individual organism, mainly including their sizes, numbers, and shapes. Karyotyping is the process by which a karyotype is disce ...
of gibbons, however, diverged in a much more rapid fashion from the common hominoid ancestor than other apes. The common ancestor of hominoids is shown to have a minimum of 24 major chromosomal rearrangements from the presumed gibbon ancestor's karyotype. Reaching the common gibbon ancestor's karyotype from today's various living species of gibbons will require up to 28 additional rearrangements. Adding up, this implies that at least 52 major chromosomal rearrangements are needed to compare the common hominoid ancestor to today's gibbons. No common specific sequence element in the independent rearrangements was found, while 46% of the gibbon-human synteny breakpoints occur in segmental duplication regions. This is an indication that these major differences in humans and gibbons could have had a common source of plasticity or change. Researchers view this unusually high rate of chromosomal rearrangement that is specific in small apes such as gibbons could potentially be due to factors that increase the rate of chromosomal breakage or factors that allow derivative chromosomes to be fixed in a homozygous state while mostly lost in other mammals. The whole genome of the gibbons in Southeast Asia was first sequenced in 2014 by the
German Primate Center The German Primate Centre (German: Deutsches Primatenzentrum, DPZ, founded in 1977) is a non-profit independent research and service institute located in Göttingen. It is a member of the Gottfried Wilhelm Leibniz Scientific Community and funded by ...
, including Christian Roos, Markus Brameier, and Lutz Walter, along with other international researchers. One of the gibbons that had its genome sequenced is a white-cheeked gibbon ('' Nomascus leucogenys'', NLE) named Asia. The team found that a jumping DNA element named LAVA transposon (also called gibbon-specific retrotransposon) is unique to the gibbon genome apart from humans and the great apes. The LAVA transposon increases mutation rate, thus is supposed to have contributed to the rapid and greater change in gibbons in comparison to their close relatives, which is critical for evolutionary development. The very high rate of chromosomal disorder and rearrangements (such as duplications, deletions or inversions of large stretches of DNA) due to the moving of this large DNA segment is one of the key features that are unique to the gibbon genome. A special feature of the LAVA transposon is that it positioned itself precisely between genes that are involved in
chromosome segregation Chromosome segregation is the process in eukaryotes by which two sister chromatids formed as a consequence of DNA replication, or paired homologous chromosomes, separate from each other and migrate to opposite poles of the nucleus. This segregation ...
and distribution during cell division, which results in a premature termination state leading to an alteration in
transcription Transcription refers to the process of converting sounds (voice, music etc.) into letters or musical notes, or producing a copy of something in another medium, including: Genetics * Transcription (biology), the copying of DNA into RNA, the fir ...
. This incorporation of the jumping gene near genes involved in chromosome replication is thought to make the rearrangement in the genome even more likely, leading to a greater diversity within the gibbon genera. In addition, some characteristic genes in the gibbon genome had gone through a positive selection and are suggested to give rise to specific anatomical features for gibbons to adapt to their new environment. One of them is ''TBX5'', which is a gene that is required for the development of the front extremities or forelimbs such as long arms. The other is ''COL1A1'', which is responsible for the development of
collagen Collagen () is the main structural protein in the extracellular matrix found in the body's various connective tissues. As the main component of connective tissue, it is the most abundant protein in mammals, making up from 25% to 35% of the whole ...
, a protein that is directly involved with the forming of connective tissues, bone, and cartilage. This gene is thought to have a role in gibbons' stronger muscles. Researchers have found a coincidence between major environmental changes in Southeast Asia about 5 Mya that caused a cyclical dynamic of expansions and contractions of their forest habitat, an instance of
radiation In physics, radiation is the emission or transmission of energy in the form of waves or particles through space or through a material medium. This includes: * ''electromagnetic radiation'', such as radio waves, microwaves, infrared, visi ...
experienced by the gibbon genera. This may have led to the development of a suite of physical characteristics, distinct from their great ape relatives, to adapt to their habitat of dense, canopy forest. These crucial findings in genetics have contributed to the use of gibbons as a genetic model for chromosome breakage and fusion, which is a type of translocation mutation. The unusually high number of structural changes in the DNA and chromosomal rearrangements could lead to problematic consequences in some species. Gibbons, however, not only seemed to be free from problems but let the change help them effectively adapt to their environment. Thus, gibbons are organisms on which genetics research could be focused to broaden the implications to human diseases related to chromosomal changes, such as cancer, including chronic myeloid leukemia.


Conservation status

Most species are either endangered or critically endangered (the sole exception being '' H. leuconedys'', which is
vulnerable Vulnerable may refer to: General * Vulnerability * Vulnerability (computing) * Vulnerable adult * Vulnerable species Music Albums * ''Vulnerable'' (Marvin Gaye album), 1997 * ''Vulnerable'' (Tricky album), 2003 * ''Vulnerable'' (The Used album) ...
), primarily due to degradation or loss of their forest habitats.A-Z Animals: Gibbon
Retrieved Feb-12-2015
On the island of Phuket in Thailand, a volunteer-based Gibbon Rehabilitation Center rescues gibbons that were kept in captivity, and are being released back into the wild. The
Kalaweit Project The Kalaweit project is a pioneer project in the conservation and protection of the gibbons of Indonesia. Origin Chanee Kalaweit The founder of the project is Aurélien Brulé (better known as "Chanee Kalaweit"), a young Frenchman born in 1979 ...
also has gibbon rehabilitation centers on Borneo and
Sumatra Sumatra is one of the Sunda Islands of western Indonesia. It is the largest island that is fully within Indonesian territory, as well as the sixth-largest island in the world at 473,481 km2 (182,812 mi.2), not including adjacent i ...
. The IUCN Species Survival Commission Primate Specialist Group announced 2015 to be the Year of the Gibbon and initiated events to be held around the world in zoos to promote awareness of the status of gibbons.


In traditional Chinese culture

Sinologist Sinology, or Chinese studies, is an academic discipline that focuses on the study of China primarily through Chinese philosophy, language, literature, culture and history and often refers to Western scholarship. Its origin "may be traced to the ex ...
Robert van Gulik Robert Hans van Gulik (, 9 August 1910 – 24 September 1967) was a Dutch orientalist, diplomat, musician (of the guqin), and writer, best known for the Judge Dee historical mysteries, the protagonist of which he borrowed from the 18th-century ...
concluded gibbons were widespread in central and southern China until at least the Song dynasty, and furthermore, based on an analysis of references to primates in Chinese poetry and other literature and their portrayal in Chinese paintings, the Chinese word ''yuán'' (猿) referred specifically to gibbons until they were extirpated throughout most of the country due to habitat destruction (around the 14th century). In modern usage, however, ''yuán'' is a generic word for ape. Early Chinese writers viewed the "noble" gibbons, gracefully moving high in the treetops, as the "gentlemen" (''jūnzǐ'', 君子) of the forest, in contrast to the greedy
macaque The macaques () constitute a genus (''Macaca'') of gregarious Old World monkeys of the subfamily Cercopithecinae. The 23 species of macaques inhabit ranges throughout Asia, North Africa, and (in one instance) Gibraltar. Macaques are principally ...
s, attracted by human food. The Taoists ascribed occult properties to gibbons, believing them to be able to live for several hundred years and to turn into humans. Gibbon figurines as old as from the fourth to third centuries BCE (the Zhou dynasty) have been found in China. Later on, gibbons became a popular subject for Chinese painters, especially during the Song dynasty and early Yuan dynasty, when Yì Yuánjí and Mùqī Fǎcháng excelled in painting these apes. From Chinese cultural influence, the Zen motif of the "gibbon grasping at the reflection of the moon in the water" became popular in
Japanese art Japanese art covers a wide range of art styles and media, including ancient pottery, sculpture, ink painting and calligraphy on silk and paper, ''ukiyo-e'' paintings and woodblock prints, ceramics, origami, and more recently manga and anime. It ...
, as well, though gibbons have never occurred naturally in Japan.Geissmann, Thomas
"Gibbon paintings in China, Japan, and Korea: Historical distribution, production rate and context"
, ''Gibbon Journal'', No. 4, May 2008. (includes color reproductions of a large number of gibbon paintings by many artists)


References


External links

* *
IUCN SSC PSG Section on Small Apes

Gibbon Conservation Center

Gibbon Network and Research Lab

Gibbon Conservation Alliance

Gibbon Rehabilitation Project
* {{Authority control Primate families Miocene primates of Asia Extant Tortonian first appearances