Rhesus Macaque
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The rhesus macaque (''Macaca mulatta''), colloquially rhesus monkey, is a species of
Old World monkey Old World monkey is the common English name for a family of primates known taxonomically as the Cercopithecidae (). Twenty-four genera and 138 species are recognized, making it the largest primate family. Old World monkey genera include baboons ...
. There are between six and nine recognised subspecies that are split between two groups, the Chinese-derived and the Indian-derived. Generally brown or grey in colour, it is in length with a tail and weighs . It is native to
South South is one of the cardinal directions or Points of the compass, compass points. The direction is the opposite of north and is perpendicular to both east and west. Etymology The word ''south'' comes from Old English ''sūþ'', from earlier Pro ...
,
Central Central is an adjective usually referring to being in the center of some place or (mathematical) object. Central may also refer to: Directions and generalised locations * Central Africa, a region in the centre of Africa continent, also known as ...
, and
Southeast Asia Southeast Asia, also spelled South East Asia and South-East Asia, and also known as Southeastern Asia, South-eastern Asia or SEA, is the geographical United Nations geoscheme for Asia#South-eastern Asia, south-eastern region of Asia, consistin ...
and has the widest geographic range of all non-human
primate Primates are a diverse order of mammals. They are divided into the strepsirrhines, which include the lemurs, galagos, and lorisids, and the haplorhines, which include the tarsiers and the simians (monkeys and apes, the latter including huma ...
s, occupying a great diversity of altitudes and a great variety of habitats, from grasslands to arid and forested areas, but also close to human settlements. Feral colonies are found in the United States, thought to be either released by humans or escapees after hurricanes destroyed zoo and wildlife park facilities. The rhesus macaque is diurnal, arboreal, and terrestrial. It is mostly herbivorous, mainly eating fruit, but will also consume seeds, roots, buds, bark, and cereals. Studies show almost 100 different plant species in its diet. Rhesus macaques are generalist omnivores, and have a highly varied and flexible diet. Text was copied from this source, which is available under
Creative Commons Attribution 4.0 International License
With an increase in anthropogenic land changes, rhesus macaques have modified their behaviours to accomodate intense and rapid environmental disturbance associated with human agriculture and urbanization resulting in proportions of their diet to be altered. It will also eat invertebrates, drink water from streams and rivers, and has specialised cheek pouches where it can temporarily store food. Like other macaques, the rhesus macaque is
gregarious Sociality is the degree to which individuals in an animal population tend to associate in social groups (gregariousness) and form cooperative societies. Sociality is a survival response to evolutionary pressures. For example, when a mother wasp ...
, with troops comprising 20-200 individuals. The social groups are
matrilineal Matrilineality is the tracing of kinship through the female line. It may also correlate with a social system in which each person is identified with their matriline – their mother's Lineage (anthropology), lineage – and which can in ...
, whereby a female's rank is decided by the rank of her mother. There has been extensive research into female
philopatry Philopatry is the tendency of an organism to stay in or habitually return to a particular area. The causes of philopatry are numerous, but natal philopatry, where animals return to their birthplace to breed, may be the most common. The term derive ...
, common in social animals, as females tend not to leave the social group. The rhesus macaque communicates with a variety of facial expressions, vocalisations, body postures, and gestures. Facial expressions are used to appease or redirect aggression, assert dominance, and threaten other individuals, and vocalisations may be made to elicit grooming, while moving, or in threatening situations. It spends most of its day feeding and resting; the rest is occupied with traveling, grooming, and playing. Due to its relatively easy upkeep, wide availability, and closeness to humans anatomically and physiologically, it has been used extensively in medical and biological research on human and animal health-related topics. It has facilitated many scientific breakthroughs including vaccines for
rabies Rabies is a viral disease that causes encephalitis in humans and other mammals. Early symptoms can include fever and tingling at the site of exposure. These symptoms are followed by one or more of the following symptoms: nausea, vomiting, vi ...
,
smallpox Smallpox was an infectious disease caused by variola virus (often called smallpox virus) which belongs to the genus Orthopoxvirus. The last naturally occurring case was diagnosed in October 1977, and the World Health Organization (WHO) c ...
, and
polio Poliomyelitis, commonly shortened to polio, is an infectious disease caused by the poliovirus. Approximately 70% of cases are asymptomatic; mild symptoms which can occur include sore throat and fever; in a proportion of cases more severe s ...
and
antiretroviral medication Antiviral drugs are a class of medication used for treating viral infections. Most antivirals target specific viruses, while a broad-spectrum antiviral is effective against a wide range of viruses. Unlike most antibiotics, antiviral drugs do n ...
to treat
HIV/AIDS Human immunodeficiency virus infection and acquired immunodeficiency syndrome (HIV/AIDS) is a spectrum of conditions caused by infection with the human immunodeficiency virus (HIV), a retrovirus. Following initial infection an individual ...
. A rhesus macaque became the first primate astronaut in 1948, but died during the flight, followed on 14 June 1949 by Albert II, who became the first primate and first mammal in space. It is listed as
Least Concern A least-concern species is a species that has been categorized by the International Union for Conservation of Nature (IUCN) as evaluated as not being a focus of species conservation because the specific species is still plentiful in the wild. T ...
in the
IUCN Red List of Threatened Species The International Union for Conservation of Nature (IUCN) Red List of Threatened Species, also known as the IUCN Red List or Red Data Book, founded in 1964, is the world's most comprehensive inventory of the global conservation status of biologi ...
in view of its wide distribution, presumed large population, and its tolerance of a broad range of
habitat In ecology, the term habitat summarises the array of resources, physical and biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species habitat can be seen as the physical ...
s.


Etymology

The name "rhesus" is reminiscent of the mythological king
Rhesus of Thrace Rhesus (; Ancient Greek: Ῥῆσος ''Rhêsos'') is a mythical Thracian king in '' Iliad'', Book X, who fought on the side of Trojans. Diomedes and Odysseus stole his team of fine horses during a night raid on the Trojan camp. Etymology His ...
, a minor character in
the Iliad The ''Iliad'' (; grc, Ἰλιάς, Iliás, ; "a poem about Ilium") is one of two major ancient Greek epic poems attributed to Homer. It is one of the oldest extant works of literature still widely read by modern audiences. As with the '' Ody ...
. However, the French naturalist
Jean-Baptiste Audebert Jean Baptiste Audebert (1759 – December 1800) was a French artist and naturalist. Life Audebert was born at Rochefort. He studied painting and drawing at Paris, and gained reputation as a miniature-painter. Employed in preparing plates ...
, who applied the name to the species, stated: "it has no meaning". The rhesus macaque is also known colloquially as the "rhesus monkey".


Taxonomy

According to Zimmermann's first description of 1780, the rhesus macaque is distributed in eastern
Afghanistan Afghanistan, officially the Islamic Emirate of Afghanistan,; prs, امارت اسلامی افغانستان is a landlocked country located at the crossroads of Central Asia and South Asia. Referred to as the Heart of Asia, it is bordere ...
,
Bangladesh Bangladesh (}, ), officially the People's Republic of Bangladesh, is a country in South Asia. It is the eighth-most populous country in the world, with a population exceeding 165 million people in an area of . Bangladesh is among the mos ...
,
Bhutan Bhutan (; dz, འབྲུག་ཡུལ་, Druk Yul ), officially the Kingdom of Bhutan,), is a landlocked country in South Asia. It is situated in the Eastern Himalayas, between China in the north and India in the south. A mountainous ...
, as far east as the
Brahmaputra Valley The Brahmaputra Valley is a region situated between hill ranges of the eastern and northeastern Himalayan range in Eastern India. The valley consists of the Western Brahmaputra Valley covering the regions of Goalpara and Kamrup; the Central ...
, Barak valley and in peninsular
India India, officially the Republic of India (Hindi: ), is a country in South Asia. It is the seventh-largest country by area, the second-most populous country, and the most populous democracy in the world. Bounded by the Indian Ocean on the so ...
,
Nepal Nepal (; ne, नेपाल ), formerly the Federal Democratic Republic of Nepal ( ne, सङ्घीय लोकतान्त्रिक गणतन्त्र नेपाल ), is a landlocked country in South Asia. It is mai ...
, and northern
Pakistan Pakistan ( ur, ), officially the Islamic Republic of Pakistan ( ur, , label=none), is a country in South Asia. It is the world's List of countries and dependencies by population, fifth-most populous country, with a population of almost 24 ...
. Today, this is known as the Indian rhesus macaque ''M. m. mulatta'', which includes the morphologically similar ''M. rhesus villosus'', described by
True True most commonly refers to truth, the state of being in congruence with fact or reality. True may also refer to: Places * True, West Virginia, an unincorporated community in the United States * True, Wisconsin, a town in the United States * ...
in 1894, from
Kashmir Kashmir () is the northernmost geographical region of the Indian subcontinent. Until the mid-19th century, the term "Kashmir" denoted only the Kashmir Valley between the Great Himalayas and the Pir Panjal Range. Today, the term encompas ...
, and ''M. m. mcmahoni'', described by Pocock in 1932 from Kootai,
Pakistan Pakistan ( ur, ), officially the Islamic Republic of Pakistan ( ur, , label=none), is a country in South Asia. It is the world's List of countries and dependencies by population, fifth-most populous country, with a population of almost 24 ...
. Several
Chinese Chinese can refer to: * Something related to China * Chinese people, people of Chinese nationality, citizenship, and/or ethnicity **''Zhonghua minzu'', the supra-ethnic concept of the Chinese nation ** List of ethnic groups in China, people of ...
subspecies In biological classification, subspecies is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics (morphology), but that can successfully interbreed. Not all species ...
of rhesus macaques were described between 1867 and 1917. The molecular differences identified among populations, however, are alone not consistent enough to conclusively define any subspecies. The Chinese subspecies can be divided as follows: * ''M. m. mulatta'' is found in western and central China, in the south of
Yunnan Yunnan , () is a landlocked Provinces of China, province in Southwest China, the southwest of the People's Republic of China. The province spans approximately and has a population of 48.3 million (as of 2018). The capital of the province is ...
, and southwest of
Guangxi Guangxi (; ; Chinese postal romanization, alternately romanized as Kwanghsi; ; za, Gvangjsih, italics=yes), officially the Guangxi Zhuang Autonomous Region (GZAR), is an Autonomous regions of China, autonomous region of the People's Republic ...
; * ''M. m. lasiota'' (
Gray Grey (more common in British English) or gray (more common in American English) is an intermediate color between black and white. It is a neutral or achromatic color, meaning literally that it is "without color", because it can be composed o ...
, 1868), the west Chinese rhesus macaque, is distributed in the west of
Sichuan Sichuan (; zh, c=, labels=no, ; zh, p=Sìchuān; alternatively romanized as Szechuan or Szechwan; formerly also referred to as "West China" or "Western China" by Protestant missions) is a province in Southwest China occupying most of the ...
, northwest of
Yunnan Yunnan , () is a landlocked Provinces of China, province in Southwest China, the southwest of the People's Republic of China. The province spans approximately and has a population of 48.3 million (as of 2018). The capital of the province is ...
, and southeast of
Qinghai Qinghai (; alternately romanized as Tsinghai, Ch'inghai), also known as Kokonor, is a landlocked province in the northwest of the People's Republic of China. It is the fourth largest province of China by area and has the third smallest po ...
; it is possibly synonymous with ''M. m. sanctijohannis'' ( R. Swinhoe, 1867), if not with ''M. m. mulatta''. * ''M. m. tcheliensis'' ( Milne-Edwards, 1870), the north Chinese rhesus macaque, lives in the north of
Henan Henan (; or ; ; alternatively Honan) is a landlocked province of China, in the central part of the country. Henan is often referred to as Zhongyuan or Zhongzhou (), which literally means "central plain" or "midland", although the name is al ...
, south of
Shanxi Shanxi (; ; formerly romanised as Shansi) is a landlocked province of the People's Republic of China and is part of the North China region. The capital and largest city of the province is Taiyuan, while its next most populated prefecture-lev ...
, and near
Beijing } Beijing ( ; ; ), alternatively romanized as Peking ( ), is the capital of the People's Republic of China. It is the center of power and development of the country. Beijing is the world's most populous national capital city, with over 21 ...
. Some consider it as the most endangered subspecies. Others consider it possibly synonymous with ''M. m. sanctijohannis'', if not with ''M. m. mulatta''. * ''M. m. vestita'' ( Milne-Edwards, 1892), the Tibetan rhesus macaque, lives in the southeast of
Tibet Tibet (; ''Böd''; ) is a region in East Asia, covering much of the Tibetan Plateau and spanning about . It is the traditional homeland of the Tibetan people. Also resident on the plateau are some other ethnic groups such as Monpa people, ...
, northwest of Yunnan ( Deqing), and perhaps including Yushu; it is possibly synonymous with ''M. m. sanctijohannis'', if not with ''M. m. mulatta''. * ''M. m. littoralis'' (
Elliot Elliot (also spelled Eliot, Elliotte, Elliott, Eliott and Elyot) is a personal name which can serve as either a surname or a given name. Although the given name has historically been given to males, females have increasingly been given the name ...
, 1909), the south Chinese rhesus macaque, lives in
Fujian Fujian (; alternately romanized as Fukien or Hokkien) is a province on the southeastern coast of China. Fujian is bordered by Zhejiang to the north, Jiangxi to the west, Guangdong to the south, and the Taiwan Strait to the east. Its capi ...
,
Zhejiang Zhejiang ( or , ; , also romanized as Chekiang) is an eastern, coastal province of the People's Republic of China. Its capital and largest city is Hangzhou, and other notable cities include Ningbo and Wenzhou. Zhejiang is bordered by Jiang ...
,
Anhui Anhui , (; formerly romanized as Anhwei) is a landlocked province of the People's Republic of China, part of the East China region. Its provincial capital and largest city is Hefei. The province is located across the basins of the Yangtze River ...
,
Jiangxi Jiangxi (; ; formerly romanized as Kiangsi or Chianghsi) is a landlocked province in the east of the People's Republic of China. Its major cities include Nanchang and Jiujiang. Spanning from the banks of the Yangtze river in the north int ...
,
Hunan Hunan (, ; ) is a landlocked province of the People's Republic of China, part of the South Central China region. Located in the middle reaches of the Yangtze watershed, it borders the province-level divisions of Hubei to the north, Jiangxi to ...
,
Hubei Hubei (; ; alternately Hupeh) is a landlocked province of the People's Republic of China, and is part of the Central China region. The name of the province means "north of the lake", referring to its position north of Dongting Lake. The prov ...
,
Guizhou Guizhou (; formerly Kweichow) is a landlocked province in the southwest region of the People's Republic of China. Its capital and largest city is Guiyang, in the center of the province. Guizhou borders the autonomous region of Guangxi to t ...
, northwest of
Guangdong Guangdong (, ), alternatively romanized as Canton or Kwangtung, is a coastal province in South China on the north shore of the South China Sea. The capital of the province is Guangzhou. With a population of 126.01 million (as of 2020) ...
, north of
Guangxi Guangxi (; ; Chinese postal romanization, alternately romanized as Kwanghsi; ; za, Gvangjsih, italics=yes), officially the Guangxi Zhuang Autonomous Region (GZAR), is an Autonomous regions of China, autonomous region of the People's Republic ...
, northeast of
Yunnan Yunnan , () is a landlocked Provinces of China, province in Southwest China, the southwest of the People's Republic of China. The province spans approximately and has a population of 48.3 million (as of 2018). The capital of the province is ...
, east of
Sichuan Sichuan (; zh, c=, labels=no, ; zh, p=Sìchuān; alternatively romanized as Szechuan or Szechwan; formerly also referred to as "West China" or "Western China" by Protestant missions) is a province in Southwest China occupying most of the ...
, and south of
Shaanxi Shaanxi (alternatively Shensi, see #Name, § Name) is a landlocked Provinces of China, province of China. Officially part of Northwest China, it borders the province-level divisions of Shanxi (NE, E), Henan (E), Hubei (SE), Chongqing (S), Sichu ...
; it is possibly synonymous with ''M. m. sanctijohannis'', if not with ''M. m. mulatta''. * ''M. m. brevicaudus'', also referred to as ''Pithecus brevicaudus'' (
Elliot Elliot (also spelled Eliot, Elliotte, Elliott, Eliott and Elyot) is a personal name which can serve as either a surname or a given name. Although the given name has historically been given to males, females have increasingly been given the name ...
, 1913), lives on the
Hainan Island Hainan (, ; ) is the smallest and southernmost province of the People's Republic of China (PRC), consisting of various islands in the South China Sea. , the largest and most populous island in China,The island of Taiwan, which is slightly ...
and Wanshan Islands in
Guangdong Guangdong (, ), alternatively romanized as Canton or Kwangtung, is a coastal province in South China on the north shore of the South China Sea. The capital of the province is Guangzhou. With a population of 126.01 million (as of 2020) ...
, and the islands near
Hong Kong Hong Kong ( (US) or (UK); , ), officially the Hong Kong Special Administrative Region of the People's Republic of China ( abbr. Hong Kong SAR or HKSAR), is a city and special administrative region of China on the eastern Pearl River Delt ...
; it may be synonymous with ''M. m. mulatta''. * ''M. m. siamica'' ( Kloss, 1917), the Indochinese rhesus macaque, is distributed in
Myanmar Myanmar, ; UK pronunciations: US pronunciations incl. . Note: Wikipedia's IPA conventions require indicating /r/ even in British English although only some British English speakers pronounce r at the end of syllables. As John C. Wells, Joh ...
, in the north of
Thailand Thailand ( ), historically known as Siam () and officially the Kingdom of Thailand, is a country in Southeast Asia, located at the centre of the Indochinese Peninsula, spanning , with a population of almost 70 million. The country is bo ...
and
Vietnam Vietnam or Viet Nam ( vi, Việt Nam, ), officially the Socialist Republic of Vietnam,., group="n" is a country in Southeast Asia, at the eastern edge of mainland Southeast Asia, with an area of and population of 96 million, making i ...
, in
Laos Laos (, ''Lāo'' )), officially the Lao People's Democratic Republic ( Lao: ສາທາລະນະລັດ ປະຊາທິປະໄຕ ປະຊາຊົນລາວ, French: République démocratique populaire lao), is a socialist ...
, and in the Chinese provinces of
Anhui Anhui , (; formerly romanized as Anhwei) is a landlocked province of the People's Republic of China, part of the East China region. Its provincial capital and largest city is Hefei. The province is located across the basins of the Yangtze River ...
, northwest
Guangxi Guangxi (; ; Chinese postal romanization, alternately romanized as Kwanghsi; ; za, Gvangjsih, italics=yes), officially the Guangxi Zhuang Autonomous Region (GZAR), is an Autonomous regions of China, autonomous region of the People's Republic ...
, Guizhou,
Hubei Hubei (; ; alternately Hupeh) is a landlocked province of the People's Republic of China, and is part of the Central China region. The name of the province means "north of the lake", referring to its position north of Dongting Lake. The prov ...
,
Hunan Hunan (, ; ) is a landlocked province of the People's Republic of China, part of the South Central China region. Located in the middle reaches of the Yangtze watershed, it borders the province-level divisions of Hubei to the north, Jiangxi to ...
, central and eastern
Sichuan Sichuan (; zh, c=, labels=no, ; zh, p=Sìchuān; alternatively romanized as Szechuan or Szechwan; formerly also referred to as "West China" or "Western China" by Protestant missions) is a province in Southwest China occupying most of the ...
, and western and south-central
Yunnan Yunnan , () is a landlocked Provinces of China, province in Southwest China, the southwest of the People's Republic of China. The province spans approximately and has a population of 48.3 million (as of 2018). The capital of the province is ...
; possibly synonymous with ''M. m. sanctijohannis'', if not with ''M. m. mulatta''.


Description

The rhesus macaque is brown or grey in color and has a pink face, which is bereft of fur. It has, on average, 50 vertebrae and a wide rib cage. Its tail averages between . Adult males measure about on average and weigh about . Females are smaller, averaging in length and in weight. The ratio of arm length to leg length is 89.6–94.3%. The rhesus macaque has a dental formula of and
bilophodont The molars or molar teeth are large, flat tooth, teeth at the back of the mouth. They are more developed in mammal, mammals. They are used primarily to comminution, grind food during mastication, chewing. The name ''molar'' derives from Latin, ...
molar teeth The molars or molar teeth are large, flat teeth at the back of the mouth. They are more developed in mammals. They are used primarily to grind food during chewing. The name ''molar'' derives from Latin, ''molaris dens'', meaning "millstone t ...
.


Distribution and Habitat

Rhesus macaques are native to
India India, officially the Republic of India (Hindi: ), is a country in South Asia. It is the seventh-largest country by area, the second-most populous country, and the most populous democracy in the world. Bounded by the Indian Ocean on the so ...
,
Bangladesh Bangladesh (}, ), officially the People's Republic of Bangladesh, is a country in South Asia. It is the eighth-most populous country in the world, with a population exceeding 165 million people in an area of . Bangladesh is among the mos ...
,
Pakistan Pakistan ( ur, ), officially the Islamic Republic of Pakistan ( ur, , label=none), is a country in South Asia. It is the world's List of countries and dependencies by population, fifth-most populous country, with a population of almost 24 ...
,
Nepal Nepal (; ne, नेपाल ), formerly the Federal Democratic Republic of Nepal ( ne, सङ्घीय लोकतान्त्रिक गणतन्त्र नेपाल ), is a landlocked country in South Asia. It is mai ...
,
Myanmar Myanmar, ; UK pronunciations: US pronunciations incl. . Note: Wikipedia's IPA conventions require indicating /r/ even in British English although only some British English speakers pronounce r at the end of syllables. As John C. Wells, Joh ...
,
Thailand Thailand ( ), historically known as Siam () and officially the Kingdom of Thailand, is a country in Southeast Asia, located at the centre of the Indochinese Peninsula, spanning , with a population of almost 70 million. The country is bo ...
,
Afghanistan Afghanistan, officially the Islamic Emirate of Afghanistan,; prs, امارت اسلامی افغانستان is a landlocked country located at the crossroads of Central Asia and South Asia. Referred to as the Heart of Asia, it is bordere ...
,
Vietnam Vietnam or Viet Nam ( vi, Việt Nam, ), officially the Socialist Republic of Vietnam,., group="n" is a country in Southeast Asia, at the eastern edge of mainland Southeast Asia, with an area of and population of 96 million, making i ...
, southern
China China, officially the People's Republic of China (PRC), is a country in East Asia. It is the world's most populous country, with a population exceeding 1.4 billion, slightly ahead of India. China spans the equivalent of five time zones and ...
, and some neighbouring areas. They have the widest geographic ranges of any non-human primate, occupying a great diversity of altitudes throughout Central, South, and Southeast Asia. Inhabiting arid, open areas, rhesus macaques may be found in grasslands, woodlands, and in mountainous regions up to in elevation. They are strong swimmers, and can swim across rivers. Rhesus macaques are noted for their tendency to move from rural to urban areas, coming to rely on handouts or refuse from humans. They adapt well to human presence, and form larger troops in human-dominated landscapes than in forests. Rhesus monkeys live in patches of forest within agricultural areas, which gives them access to agroecosystem habitats and makes them at ease in navigating through them. The southern and the northern distributional limits for rhesus and
bonnet macaque The bonnet macaque (''Macaca radiata''), also known as zati,Chambers English Dictionary is a species of macaque endemic to southern India. Its distribution is limited by the Indian Ocean on three sides and the Godavari and Tapti Rivers, along wi ...
s, respectively, currently run parallel to each other in the western part of India, are separated by a large gap in the center, and converge on the eastern coast of the peninsula to form a distribution overlap zone. This overlap region is characterized by the presence of mixed-species troops, with pure troops of both species sometimes occurring even in close proximity to one another. The range extension of rhesus macaque – a natural process in some areas, and a direct consequence of introduction by humans in other regions – poses grave implications for the endemic and declining populations of bonnet macaques in southern India. Kumar et al (2013) provides a summary of population distribution and habitat in India. It states that there were sightings of rhesus macaques in all surveyed habitats except semi-evergreen forests.


Fossil Record

Fossilized isolated teeth and
mandible In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower teeth in place. The mandible sits beneath the maxilla. It is the only movable bone ...
fragments from
Tianyuan Cave Tianyuan Cave () is near Beijing, where Tianyuan man, one of the earliest modern humans, was found. The remains in the Tianyuan Cave have ancestral relations "to many present-day Asians and Native Americans". See also * Niah Caves * Fuyan Cave ...
and a juvenile
maxilla The maxilla (plural: ''maxillae'' ) in vertebrates is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. The t ...
from Wanglaopu Cave near
Zhoukoudian Zhoukoudian Area () is a town and an area located on the east Fangshan District, Beijing, China. It borders Nanjiao and Fozizhuang Townships to its north, Xiangyang, Chengguan and Yingfeng Subdistricts to its east, Shilou and Hangcunhe Towns t ...
represent the first recognized occurrence of rhesus macaque fossils in the far north of China, and thus the population of rhesus macaques which lived around
Beijing } Beijing ( ; ; ), alternatively romanized as Peking ( ), is the capital of the People's Republic of China. It is the center of power and development of the country. Beijing is the world's most populous national capital city, with over 21 ...
decades ago is believed to have originated from
Pleistocene The Pleistocene ( , often referred to as the ''Ice age'') is the geological Epoch (geology), epoch that lasted from about 2,580,000 to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was fina ...
ancestors rather than being human-introduced. Fossil
mandible In anatomy, the mandible, lower jaw or jawbone is the largest, strongest and lowest bone in the human facial skeleton. It forms the lower jaw and holds the lower teeth in place. The mandible sits beneath the maxilla. It is the only movable bone ...
fragments from the
Taedong River The Taedong River (Chosŏn'gŭl: ) is a large river in North Korea. The river rises in the Rangrim Mountains of the country's north where it then flows southwest into Korea Bay at Namp'o.Suh, Dae-Sook (1987) "North Korea in 1986: Strengthenin ...
Basin around
Pyongyang Pyongyang (, , ) is the capital and largest city of North Korea, where it is known as the "Capital of the Revolution". Pyongyang is located on the Taedong River about upstream from its mouth on the Yellow Sea. According to the 2008 populatio ...
, North Korea, have also been assigned to this species.


Feral Colonies

Rhesus macaques have also been introduced to other areas, such as the United States, and become feral. Around the spring of 1938, a colony of rhesus macaques was released in and around Silver Springs in
Florida Florida is a state located in the Southeastern region of the United States. Florida is bordered to the west by the Gulf of Mexico, to the northwest by Alabama, to the north by Georgia, to the east by the Bahamas and Atlantic Ocean, and to ...
by a tour boat operator known locally as "Colonel Tooey" to enhance his "Jungle Cruise". A traditional story that the monkeys were released for scenery enhancement in the
Tarzan Tarzan (John Clayton II, Viscount Greystoke) is a fictional character, an archetypal feral child raised in the African jungle by the Mangani great apes; he later experiences civilization, only to reject it and return to the wild as a heroic adv ...
movies that were filmed at that location is false, as the only Tarzan movie filmed in the area, 1939's ''
Tarzan Finds a Son! ''Tarzan Finds a Son!'' is a 1939 Tarzan film based on the character created by Edgar Rice Burroughs. It was the fourth in the MGM ''Tarzan'' series to feature Johnny Weissmuller as the "King of the Apes" and the fourth of six films in which he ...
'', does not contain rhesus macaques. Various colonies of rhesus macaque are speculated to be the result of
zoos A zoo (short for zoological garden; also called an animal park or menagerie) is a facility in which animals are kept within enclosures for public exhibition and often bred for conservation purposes. The term ''zoological garden'' refers to zool ...
and wildlife parks destroyed in hurricanes, most notably
Hurricane Andrew Hurricane Andrew was a very powerful and destructive Category 5 Atlantic hurricane that struck the Bahamas, Florida, and Louisiana in August 1992. It is the most destructive hurricane to ever hit Florida in terms of structures damaged ...
. A 2020 estimate put the number at 550–600 rhesus macaques living in the state; officials have caught more than 1000 of the monkeys in the past decade. Most of the captured monkeys tested positive for
herpes B virus B-virus (''Macacine alphaherpesvirus 1''; McHV-1; formerly ''Macacine herpesvirus 1'', ''Cercopithecine herpesvirus 1'', CHV-1), ''Herpesvirus simiae'', or ''Herpes virus B'' is the ''Simplexvirus'' infecting macaque monkeys. B virus is very sim ...
, which leads wildlife officials to consider the animals a public health hazard. Feral colonies have also resulted from research activities. There is a colony of rhesus macaques on Morgan Island, one of the
Sea Islands The Sea Islands are a chain of tidal and barrier islands on the Atlantic Ocean coast of the Southeastern United States. Numbering over 100, they are located between the mouths of the Santee and St. Johns Rivers along the coast of South Carolina, ...
in the
South Carolina Lowcountry The Lowcountry (sometimes Low Country or just low country) is a geographic and cultural region along South Carolina's coast, including the Sea Islands. The region includes significant salt marshes and other coastal waterways, making it an impor ...
. They were imported in the 1970s for use in the local labs. Another research colony was established by the Caribbean Primate Research Center of the
University of Puerto Rico The University of Puerto Rico ( es, Universidad de Puerto Rico, UPR) is the main public university system in the U.S. Commonwealth of Puerto Rico. It is a government-owned corporation with 11 campuses and approximately 58,000 students and 5,3 ...
on the island of
Cayo Santiago Cayo Santiago, also known as Santiago Island, Isla de los monos (or Island of the monkeys), is located at , to the east of ''Punta Santiago'', Humacao, Puerto Rico. Geography The island measures approximately , north–south and east–west, ...
, off of
Puerto Rico Puerto Rico (; abbreviated PR; tnq, Boriken, ''Borinquen''), officially the Commonwealth of Puerto Rico ( es, link=yes, Estado Libre Asociado de Puerto Rico, lit=Free Associated State of Puerto Rico), is a Caribbean island and Unincorporated ...
. There are no predators on the island, and humans are not permitted to land, except as part of the research program. Another Puerto Rico research colony was released into the
Desecheo National Wildlife Refuge The Desecheo National Wildlife Refuge (Spanish: ''Refugio Nacional de Vida Silvestre de Desecheo'') is a National Wildlife Refuge in Puerto Rico. It is part of the Caribbean Islands National Wildlife Refuge Complex. The island of Desecheo is ...
in 1966. they are continuing to do ecological harm, damage crops amounting to $300,000/year and cost $1,000,000/year to manage.


Ecology and Behavior

Rhesus macaques are
diurnal animal Diurnality is a form of plant and animal behavior characterized by activity during daytime, with a period of sleeping or other inactivity at night. The common adjective used for daytime activity is "diurnal". The timing of activity by an animal d ...
s, and both
arboreal Arboreal locomotion is the Animal locomotion, locomotion of animals in trees. In habitats in which trees are present, animals have evolved to move in them. Some animals may scale trees only occasionally, but others are exclusively arboreal. Th ...
and
terrestrial Terrestrial refers to things related to land or the planet Earth. Terrestrial may also refer to: * Terrestrial animal, an animal that lives on land opposed to living in water, or sometimes an animal that lives on or near the ground, as opposed to ...
. They are
quadruped Quadrupedalism is a form of locomotion where four limbs are used to bear weight and move around. An animal or machine that usually maintains a four-legged posture and moves using all four limbs is said to be a quadruped (from Latin ''quattuor' ...
al and, when on the ground, they walk digitigrade and plantigrade. They are mostly
herbivorous A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpart ...
, feeding mainly on
fruit In botany, a fruit is the seed-bearing structure in flowering plants that is formed from the ovary after flowering. Fruits are the means by which flowering plants (also known as angiosperms) disseminate their seeds. Edible fruits in particu ...
, but also eating
seed A seed is an embryonic plant enclosed in a protective outer covering, along with a food reserve. The formation of the seed is a part of the process of reproduction in seed plants, the spermatophytes, including the gymnosperm and angiospe ...
s,
root In vascular plants, the roots are the organs of a plant that are modified to provide anchorage for the plant and take in water and nutrients into the plant body, which allows plants to grow taller and faster. They are most often below the sur ...
s, buds, bark, and
cereal A cereal is any Poaceae, grass cultivated for the edible components of its grain (botanically, a type of fruit called a caryopsis), composed of the endosperm, Cereal germ, germ, and bran. Cereal Grain, grain crops are grown in greater quantit ...
s. They are estimated to consume around 99 different plant species in 46 families. During the
monsoon A monsoon () is traditionally a seasonal reversing wind accompanied by corresponding changes in precipitation but is now used to describe seasonal changes in atmospheric circulation and precipitation associated with annual latitudinal oscil ...
season, they get much of their water from ripe and succulent fruit. Macaques living far from water sources lick dewdrops from leaves and drink rainwater accumulated in tree hollows. They have also been observed eating
termite Termites are small insects that live in colonies and have distinct castes (eusocial) and feed on wood or other dead plant matter. Termites comprise the infraorder Isoptera, or alternatively the epifamily Termitoidae, within the order Blattode ...
s, grasshoppers,
ant Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from vespoid wasp ancestors in the Cretaceous period. More than 13,800 of an estimated total of 22 ...
s, and
beetle Beetles are insects that form the order Coleoptera (), in the superorder Endopterygota. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 400,000 describ ...
s. When food is abundant, they are distributed in patches, and forage throughout the day in their home ranges. They drink water when foraging, and gather around streams and rivers. Rhesus macaques have specialized pouch-like cheeks, allowing them to temporarily hoard their food. In psychological research, rhesus macaques have demonstrated a variety of complex cognitive abilities, including the ability to make same-different judgments, understand simple rules, and monitor their own mental states. They have even been shown to demonstrate
self-agency Self-agency, also known as the phenomenal will, is the sense that actions are self-generated. Scientist Benjamin Libet was the first to study it, concluding that brain activity predicts the action before one even has conscious awareness of his or h ...
, an important type of self-awareness. In 2014, onlookers at a train station in Kanpur, India, documented a rhesus monkey, knocked unconscious by overhead power lines, that was revived by another rhesus that systematically administered a series of resuscitative actions.


Group Structure

Like other macaques, rhesus troops comprise a mixture of 20–200 males and females. Females may outnumber the males by a ratio of 4:1. Males and females both have separate hierarchies. Female
philopatry Philopatry is the tendency of an organism to stay in or habitually return to a particular area. The causes of philopatry are numerous, but natal philopatry, where animals return to their birthplace to breed, may be the most common. The term derive ...
, common among social mammals, has been extensively studied in rhesus macaques. Females tend not to leave the social group, and have highly stable
matrilineal Matrilineality is the tracing of kinship through the female line. It may also correlate with a social system in which each person is identified with their matriline – their mother's Lineage (anthropology), lineage – and which can in ...
hierarchies in which a female's rank is dependent on the rank of her mother. In addition, a single group may have multiple matrilineal lines existing in a hierarchy, and a female outranks any unrelated females that rank lower than her mother. Rhesus macaques are unusual in that the youngest females tend to outrank their older sisters. This is likely because young females are more fit and fertile. Mothers seem to prevent the older daughters from forming coalitions against her. The youngest daughter is the most dependent on the mother, and would have nothing to gain from helping her siblings in overthrowing their mother. Since each daughter had a high rank in her early years, rebelling against her mother is discouraged. Juvenile male macaques also exist in matrilineal lines, but once they reach four to five years of age, they are driven out of their natal groups by the dominant male. Thus, adult males gain dominance by age and experience. In the group, macaques position themselves based on rank. The "central male subgroup" contains the two or three oldest and most dominant males which are codominant, along with females, their infants, and juveniles. This subgroup occupies the center of the group and determines the movements, foraging, and other routines. The females of this subgroup are also the most dominant of the entire group. The farther to the periphery a subgroup is, the less dominant it is. Subgroups on the periphery of the central group are run by one dominant male, of a rank lower than the central males, and he maintains order in the group, and communicates messages between the central and peripheral males. A subgroup of subordinate, often subadult, males occupy the very edge of the groups, and have the responsibility of communicating with other macaque groups and making alarm calls. Rhesus social behaviour has been described as despotic, in that high-ranking individuals often show little tolerance, and frequently become aggressive towards non-kin. Top-ranking female rhesus monkeys are known to sexually coerce unreceptive males and also physically injure them, biting off digits and damaging their genitals. Rhesus macaques have been observed engaging in interspecies grooming with
Hanuman langur Gray langurs, also called Hanuman langurs and Hanuman monkeys, are Old World monkeys native to the Indian subcontinent constituting the genus ''Semnopithecus''. Traditionally only one species ''Semnopithecus entellus'' was recognized, but since a ...
s and with Sambar deer.


Communication

Rhesus macaques interact using a variety of facial expressions, vocalizations, body postures, and gestures. Perhaps the most common facial expression the macaque makes is the "silent bared teeth" face. This is made between individuals of different social ranks, with the lower-ranking one giving the expression to its superior. A less-dominant individual also makes a "fear grimace", accompanied by a scream, to appease or redirect aggression. Another submissive behavior is the "present rump", where an individual raises its tail and exposes its genitals to the dominant one. A dominant individual threatens another individual by standing quadrupedally and making a silent "open mouth stare" accompanied by the tail sticking straight. During movements, macaques make coos and grunts. These are also made during affiliative interactions, and approaches before grooming. When they find rare food of high quality, macaques emit warbles, harmonic arches, or chirps. When in threatening situations, macaques emit a single loud, high-pitched sound called a shrill bark. Screeches, screams, squeaks, pant-threats, growls, and barks are used during aggressive interactions. Infants " gecker" to attract their mother's attention.


Reproduction

Adult male macaques try to maximize their reproductive success by entering into
sex Sex is the trait that determines whether a sexually reproducing animal or plant produces male or female gametes. Male plants and animals produce smaller mobile gametes (spermatozoa, sperm, pollen), while females produce larger ones ( ova, of ...
with females both in and outside the breeding period. Females prefer to mate with males that are not familiar to them. Outsider males who are not members of the female's own troop are preferred over higher-ranking males. Outside of the consortship period, males and females return the prior behavior of not exhibiting preferential treatment or any special relationship. The breeding period can last up to eleven days, and a female usually mates with numerous males during that time. Male rhesus macaques have been observed to fight for access to sexually receptive females and they suffer more wounds during the mating season. Female macaques first breed when they are four years old and reach
menopause Menopause, also known as the climacteric, is the time in women's lives when menstrual periods stop permanently, and they are no longer able to bear children. Menopause usually occurs between the age of 47 and 54. Medical professionals often ...
at around twenty-five years of age. Male macaques generally play no role in raising the young but do have peaceful relationships with the offspring of their consort pairs. Manson and Parry found that free-ranging rhesus macaques avoid inbreeding. Adult females were never observed to copulate with males of their own matrilineage during their fertile periods. Mothers with one or more immature daughters in addition to their infants are in contact with their infants less than those with no older immature daughters, because the mothers may pass the parenting responsibilities to their daughters. High-ranking mothers with older immature daughters also reject their infants significantly more than those without older daughters and tend to begin mating earlier in the mating season than expected based on their dates of
parturition Birth is the act or process of bearing or bringing forth offspring, also referred to in technical contexts as parturition. In mammals, the process is initiated by hormones which cause the muscular walls of the uterus to contract, expelling the f ...
the preceding birth season. Infants farther from the center of the groups are more vulnerable to
infanticide Infanticide (or infant homicide) is the intentional killing of infants or offspring. Infanticide was a widespread practice throughout human history that was mainly used to dispose of unwanted children, its main purpose is the prevention of reso ...
from outside groups. Some mothers abuse their infants, which is believed to be the result of controlling
parenting styles A parenting style is a psychological construct representing standard strategies that parents use in their child rearing. The quality of parenting can be more essential than the quantity of time spent with the child. For instance, the parent may be ...
.


Self-awareness

In several
experiments An experiment is a procedure carried out to support or refute a hypothesis, or determine the efficacy or likelihood of something previously untried. Experiments provide insight into cause-and-effect by demonstrating what outcome occurs when ...
giving mirrors to rhesus monkeys, they looked into the mirrors and groomed themselves, as well as flexed various muscle groups. This behaviour indicates that they recognised and were aware of themselves.


Human - Rhesus Conflict

The Macaque–human relationships is complex and culturally specific, ranging from relatively peaceful coexistence to extreme levels of conflict. The relationship between rhesus macaques and humans is in constant change, with conflict being shaped by historic changes in social and cultural practices.  The changing perceptions of nature and human-nature relationships is influenced by larger political-economic decisions. When looking at conflict between humans and rhesus macaques there lacks an integrative approach that draws upon multiple fields to provide a more holistic understanding of the emergence and evolution of this conflict. Conflicts can be as a result of  rapidly changing agricultural practices, increasing infrastructure to support urbanisation, and emerging economic activities (e.g., tourism, food processing etc.) requiring more clearing of land including forests, and rising numbers of rhesus macaques. The issue is multi-dimensional and has a direct connection to overall economic policy – more specifically the relationship among agricultural, forest, and land use policies.  Deeply understanding factors relating to conflict is all the more critical in an uncertain and unpredictable future of climate change that is likely to increase the vulnerability of fragile mountain ecosystems and marginal communities. Conflict between Rhesus Macaques and humans is at all time high, with areas once forested habitat being converted to industrial agriculture. Specifically looking at Nepal, this process has increased urban infrastructure such as housing and roads that increasingly fragment forest ecosystems. The expansion of monocultures, increased forest fragmentation, degradation of natural habitats, and changing agricultural practices have led to a significant increase in the frequency of human-macaque conflict.Crop raiding is one of the biggest visible effects of Human-Rhesus conflict occurring where Rhesus Macaque feed on growing crops that directly effected harvest size, and crop health with corn, and rice. The estimated financial cost to individual farmer households of macaque corn and rice raiding is approximately US$ 14.9 or 4.2% of their yearly income.This has resulted in farmers and other members of the population viewing macaques inhabiting agricultural landscapes as serious crop pests.Nepal is a significant study area with almost 44% of Nepal's land area containing suitable habitat for rhesus macaques but only having 8% of such suitable area being protected national parks. As well the rating of Rhesus macaques as the top ten crop-raiding wildlife species in Nepal adds to such negative perception. Studying crop raiding behaviour is essential to developing effective strategies to manage human-macaque conflict while promoting both primate conservation and the economic well-being of the local community. It is stated that the human-macaque conflict is one of the most critical challenges faced by wildlife managers. Suggestions to mitigate conflict include "prioritizing forest restoration programs, strategic management plans designed to connect isolated forest fragments with high rhesus macaque population densities, creating government programs that compensate farmers for income lost due to crop-raiding, and educational outreach that informs local villagers of the importance of conservation and protecting biodiversity". Mitigation strategies offers the most effective solutions to reduce conflict occurring between rhesus macaques and humans in Nepal. India is another country that is seeing the rise of human - macaque conflict. Macaque-Human conflict particularly occurs in the twin hill-states of Uttarakhand and Himachal Pradesh with such conflict being a source of contentious debate in political scenarios, resentment and polarization amongst agriculturalists and wildlife conservationists. In India crop raiding by rhesus macaques has been identified as the main cause of conflict. In urban areas, rhesus macaques damage property and injure people in house raids in order to access food and provisions. Whereas in agricultural areas, they cause financial losses to farmers due to crop depredation. The estimated extent of crop damages in Himachal Pradesh ranges from 10–100% to 40–80% of all crop losses. The financial implications of such damage is estimated at approximately USD$ 200,000 in agriculture and USD$ 150,000 in horticulture. Quantification of crop and financial loses is challenging with a potential misrepresentation due to farmer perspectives where perception of perceived losses are potentially higher, than actual losses. This has led to harsh actions against rhesus macaque communities. Another factor in rhesus perception includes economic status, farmer economic stability, cultural attitudes towards the given species and the frequency and intensity of wildlife conflicts. All of the above have resulted in changed in conservation and management with legal rhesus macaque culling issued in 2010. Human wildlife conflict is also occurring in China, specifically in the area of Longyang District, Baoshan City, Yunnan Province. Sources have established that the peak period of conflict occurred from August-October when wildlife overlapped with humans severely due to the high natural productivity stemming from the warm and humid climate. In this research factors associated with accessibility and availability of food and shelter have been regarded as key drivers of human - macaque wildlife with an overall increase between the years of 2012 and 2021. One key factor of conflict that directly effects the human macaque relationship is visibility. Visibility of rhesus macaques in agroecosystem dominated areas largely impacts conflict between humans and rhesus macaques. The conspicuous presence of rhesus macaques in and around farms results in farmers believing that macaques cause heavy crop depredations which, in turn, have led to negative perceptions and actions against the species. Whereas visibility in urban areas can result in a positive relationship, areas include around temples, and tourist areas where their dietary needs are largely met by food provisioning.   Towards the end of March 2018, it was reported that a monkey had entered a house in the village of Talabasta, Indian state of
Odisha Odisha (English: , ), formerly Orissa ( the official name until 2011), is an Indian state located in Eastern India. It is the 8th largest state by area, and the 11th largest by population. The state has the third largest population of ...
, and kidnapped a baby. The baby was later found dead in a well. Though monkeys are known to attack people, enter homes, and damage property, this reported behaviour was unusual.


Population Management Tools

Managing conflict between humans and rhesus macaque is a difficult challenge. As mentioned previously there are many factors that go into why conflict occurs. This nuanced relationship requires thoughtfulness in management practices. Behaviour and population management are the two main areas of management that humans will look into to try and minimize conflict, protect wildlife, and promote co-existence.   When looking at and altering behaviour, crop raiding is the potentially the most significant behaviour change that is crucial in reducing conflict rates. One example is the implementation of guards in agricultural setting to chase off intruding monkeys using dogs slingshot, and firecrackers. This method is no-lethal and can alter behavioural patterns of crop raiding monkeys. Another strategy that farmers can employ is to plant alternative, buffer crops which are unattractive to monkeys in high-conflict zones such as along the edges of macaque habitat. In urban settings planting food trees within city periphery and country parks aim to discourage macaques from entering nearby residential areas for food. Better establishing tourism and urban behaviour in areas that have population of rhesus macaques as means to facilitate better relationship. In areas of tourism human behaviour is necessary to prevent conflict. One method of this is to introduce public education programs as well as  restrict visitors to specific viewing platforms, with the goal to minimize physical proximity. An important aspect is enforcing no feed regulations that only allow provisioning to be performed by trained staff at scheduled times. Regulating visitor behaviours that provoke aggressive responses from macaques, including noise regulation greatly benefits conflict reduction. Replacing food conditioned behaviours established by human visitors and further human education will greatly aid in returning co-existence between rhesus macaques and humans. A method of population management is translocation. Translocation of problem macaques in urban rhesus communities in India has been employed as a non-lethal solution to human–macaque conflicts. Translocation can be seen as a short term fix due to the fact that they have the potential to return, and other rhesus macaques populations may take their place. As well translocation can be inappropriate when there is a lack of suitable habitat to move animals because of anthropogenic habitat modification. Before translocation occurs there must be a cost benefit appraisal of relative costs should be done to quantify the resources it will take. An in depth understanding of issues prior to translocation is vital for positive effects to occur. Recognizing landscape health and productivity is the first step before making management decisions. Another tool of population management is found in sterilisation and/or contraceptive programmes that represent an alternative management practice. Fertility control looks to be a feasible management tool for reducing human–macaque conflict because it avoids the extermination of the animals and avoids costs and problems associated with translocation. Although there is potential of sterilization and general fertility control to be positive there is limited research and understanding of the long term effects of sterilization programs and its effectivity.


In science

The rhesus macaque is well known to science. Due to its relatively easy upkeep in captivity, wide availability, and closeness to humans anatomically and physiologically, it has been used extensively in medical and biological research on human and animal health-related topics. It has given its name to the
Rh factor The Rh blood group system is a human blood group system. It contains proteins on the surface of red blood cells. After the ABO blood group system, it is the most likely to be involved in transfusion reactions. The Rh blood group system consists ...
, one of the elements of a person's
blood group A blood type (also known as a blood group) is a classification of blood, based on the presence and absence of antibodies and inherited antigenic substances on the surface of red blood cells (RBCs). These antigens may be proteins, carbohydrates ...
, by the discoverers of the factor,
Karl Landsteiner Karl Landsteiner (; 14 June 1868 – 26 June 1943) was an Austrian-born American biologist, physician, and immunologist. He distinguished the main blood groups in 1900, having developed the modern system of classification of blood groups from ...
and Alexander Wiener. The rhesus macaque was also used in the well-known experiments on maternal deprivation carried out in the 1950s by
controversial Controversy is a state of prolonged public dispute or debate, usually concerning a matter of conflicting opinion or point of view. The word was coined from the Latin ''controversia'', as a composite of ''controversus'' – "turned in an opposite ...
comparative psychologist Comparative psychology refers to the scientific study of the behavior and mental processes of non-human animals, especially as these relate to the phylogenetic history, adaptive significance, and development of behavior. Research in this area addr ...
Harry Harlow Harry Frederick Harlow (October 31, 1905 – December 6, 1981) was an American psychologist best known for his maternal-separation, dependency needs, and social isolation experiments on rhesus monkeys, which manifested the importance of caregi ...
. Other medical breakthroughs facilitated by the use of the rhesus macaque include: * development of the
rabies Rabies is a viral disease that causes encephalitis in humans and other mammals. Early symptoms can include fever and tingling at the site of exposure. These symptoms are followed by one or more of the following symptoms: nausea, vomiting, vi ...
,
smallpox Smallpox was an infectious disease caused by variola virus (often called smallpox virus) which belongs to the genus Orthopoxvirus. The last naturally occurring case was diagnosed in October 1977, and the World Health Organization (WHO) c ...
, and
polio Poliomyelitis, commonly shortened to polio, is an infectious disease caused by the poliovirus. Approximately 70% of cases are asymptomatic; mild symptoms which can occur include sore throat and fever; in a proportion of cases more severe s ...
vaccines A vaccine is a biological preparation that provides active acquired immunity The adaptive immune system, also known as the acquired immune system, is a subsystem of the immune system that is composed of specialized, systemic cells and pro ...
* creation of drugs to manage
HIV/AIDS Human immunodeficiency virus infection and acquired immunodeficiency syndrome (HIV/AIDS) is a spectrum of conditions caused by infection with the human immunodeficiency virus (HIV), a retrovirus. Following initial infection an individual ...
* understanding of the female reproductive cycle and development of the embryo and the propagation of
embryonic stem cells Embryonic stem cells (ESCs) are pluripotent stem cells derived from the inner cell mass of a blastocyst, an early-stage pre- implantation embryo. Human embryos reach the blastocyst stage 4–5 days post fertilization, at which time they consist ...
. The
U.S. Army The United States Army (USA) is the land service branch of the United States Armed Forces. It is one of the eight U.S. uniformed services, and is designated as the Army of the United States in the U.S. Constitution.Article II, section 2, cl ...
, the
U.S. Air Force The United States Air Force (USAF) is the air service branch of the United States Armed Forces, and is one of the eight uniformed services of the United States. Originally created on 1 August 1907, as a part of the United States Army Sign ...
, and
NASA The National Aeronautics and Space Administration (NASA ) is an independent agency of the US federal government responsible for the civil space program, aeronautics research, and space research. NASA was established in 1958, succeeding t ...
launched rhesus macaques into outer space during the 1950s and 1960s, and the Soviet/Russian space program launched them into space as recently as 1997 on the Bion missions. Albert II became the first primate and first mammal in space during a U.S.
V-2 rocket The V-2 (german: Vergeltungswaffe 2, lit=Retaliation Weapon 2), with the technical name ''Aggregat 4'' (A-4), was the world’s first long-range guided ballistic missile. The missile, powered by a liquid-propellant rocket engine, was develop ...
suborbital flight on 14 June 1949, and died on impact when a parachute failed. Another rhesus monkey, Able, was launched on a suborbital spaceflight in 1959, and was among the first living beings (along with
Miss Baker Miss Baker (1957 – November 29, 1984) was a squirrel monkey who in 1959 became, along with female rhesus macaque Able, one of the first two animals launched into space by the United States who safely returned. Previous animal flights Previous ...
, a
squirrel monkey Squirrel monkeys are New World monkeys of the genus ''Saimiri''. ''Saimiri'' is the only genus in the subfamily Saimirinae. The name of the genus is of Tupi origin (''sai-mirím'' or ''çai-mbirín'', with ''sai'' meaning 'monkey' and ''mirím'' ...
on the same mission) to travel in space and return alive. On 25 October 1999, the rhesus macaque became the first
cloned Cloning is the process of producing individual organisms with identical or virtually identical DNA, either by natural or artificial means. In nature, some organisms produce clones through asexual reproduction. In the field of biotechnology, c ...
primate Primates are a diverse order of mammals. They are divided into the strepsirrhines, which include the lemurs, galagos, and lorisids, and the haplorhines, which include the tarsiers and the simians (monkeys and apes, the latter including huma ...
with the birth of
Tetra Terrestrial Trunked Radio (TETRA; formerly known as Trans-European Trunked Radio), a European standard for a trunked radio system, is a professional mobile radio and two-way transceiver specification. TETRA was specifically designed for use by ...
. January 2001 had the birth of
ANDi Andi or ANDI may refer to: People and fictional characters * Andy (given name), including people and fictional characters with the name Andi * Andi people, an ethnic group of Dagestan, Russia Places * Andi, Guizhou, a town in Jinsha County, Gu ...
, the first
transgenic A transgene is a gene that has been transferred naturally, or by any of a number of genetic engineering techniques, from one organism to another. The introduction of a transgene, in a process known as transgenesis, has the potential to change the ...
primate; ANDi carries foreign
gene In biology, the word gene (from , ; "...Wilhelm Johannsen coined the word gene to describe the Mendelian units of heredity..." meaning ''generation'' or ''birth'' or ''gender'') can have several different meanings. The Mendelian gene is a ba ...
s originally from a
jellyfish Jellyfish and sea jellies are the informal common names given to the medusa-phase of certain gelatinous members of the subphylum Medusozoa, a major part of the phylum Cnidaria. Jellyfish are mainly free-swimming marine animals with umbrella- ...
. Though most studies of the rhesus macaque are from various locations in northern India, some knowledge of the natural behavior of the species comes from studies carried out on a colony established by the Caribbean Primate Research Center of the
University of Puerto Rico The University of Puerto Rico ( es, Universidad de Puerto Rico, UPR) is the main public university system in the U.S. Commonwealth of Puerto Rico. It is a government-owned corporation with 11 campuses and approximately 58,000 students and 5,3 ...
on the island of
Cayo Santiago Cayo Santiago, also known as Santiago Island, Isla de los monos (or Island of the monkeys), is located at , to the east of ''Punta Santiago'', Humacao, Puerto Rico. Geography The island measures approximately , north–south and east–west, ...
, off
Puerto Rico Puerto Rico (; abbreviated PR; tnq, Boriken, ''Borinquen''), officially the Commonwealth of Puerto Rico ( es, link=yes, Estado Libre Asociado de Puerto Rico, lit=Free Associated State of Puerto Rico), is a Caribbean island and Unincorporated ...
. No predators are on the island, and humans are not permitted to land except as part of the research programmes. The colony is provisioned to some extent, but about half of its food comes from natural foraging. Rhesus macaques, like many macaques, carry the
herpes B virus B-virus (''Macacine alphaherpesvirus 1''; McHV-1; formerly ''Macacine herpesvirus 1'', ''Cercopithecine herpesvirus 1'', CHV-1), ''Herpesvirus simiae'', or ''Herpes virus B'' is the ''Simplexvirus'' infecting macaque monkeys. B virus is very sim ...
. This virus does not typically harm the monkey, but is very dangerous to humans in the rare event that it jumps species, for example in the 1997 death of
Yerkes National Primate Research Center The Emory National Primate Research Center (formerly known as Yerkes National Primate Research Center) located in Atlanta, Georgia, owned by Emory University, is a center of biomedical and behavioral research, is dedicated to improving human and a ...
researcher Elizabeth Griffin.


Genome Sequencing

Work on the
genome In the fields of molecular biology and genetics, a genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding ge ...
of the rhesus macaque was completed in 2007, making the species the second nonhuman
primate Primates are a diverse order of mammals. They are divided into the strepsirrhines, which include the lemurs, galagos, and lorisids, and the haplorhines, which include the tarsiers and the simians (monkeys and apes, the latter including huma ...
whose genome was sequenced. Humans and macaques apparently share about 93% of their
DNA sequence DNA sequencing is the process of determining the nucleic acid sequence – the order of nucleotides in DNA. It includes any method or technology that is used to determine the order of the four bases: adenine, guanine, cytosine, and thymine. Th ...
and shared a
common ancestor Common descent is a concept in evolutionary biology applicable when one species is the ancestor of two or more species later in time. All living beings are in fact descendants of a unique ancestor commonly referred to as the last universal comm ...
roughly 25 million years ago. The rhesus macaque has 21 pairs of chromosomes. Comparison of rhesus macaques, chimpanzees, and humans revealed the structure of ancestral primate genomes, positive selection pressure and lineage-specific expansions, and contractions of gene families. "The goal is to reconstruct the history of every gene in the human genome," said Evan Eichler, University of Washington, Seattle. DNA from different branches of the primate tree will allow us "to trace back the evolutionary changes that occurred at various time points, leading from the common ancestors of the primate clade to ''Homo sapiens''," said
Bruce Lahn Bruce Lahn is a Chinese-born American geneticist. Lahn came to the U.S. from China to continue his education in the late 1980s. He is the William B. Graham professor of Human Genetics at the University of Chicago. He is also the founder of the Cen ...
, University of Chicago. After the human and chimpanzee genomes were sequenced and compared, it was usually impossible to tell whether differences were the result of the human or chimpanzee gene changing from the common ancestor. After the rhesus macaque genome was sequenced, three genes could be compared. If two genes were the same, they were presumed to be the original gene. The chimpanzee and human genome diverged 6 million years ago. They have 98% identity and many conserved regulatory regions. Comparing the macaque and human genomes, further identified evolutionary pressure and gene function. Like the chimpanzee, changes were on the level of gene rearrangements rather than single mutations. Frequent insertions, deletions, changes in the order and number of genes, and segmental duplications near gaps, centromeres and telomeres occurred. So, macaque, chimpanzee, and human chromosomes are mosaics of each other. Some normal gene sequences in healthy macaques and chimpanzees cause profound disease in humans. For example, the normal sequence of phenylalanine hydroxylase in macaques and chimpanzees is the mutated sequence responsible for
phenylketonuria Phenylketonuria (PKU) is an inborn error of metabolism that results in decreased metabolism of the amino acid phenylalanine. Untreated PKU can lead to intellectual disability, seizures, behavioral problems, and mental disorders. It may also re ...
in humans. So, humans must have been under evolutionary pressure to adopt a different mechanism. Some gene families are conserved or under evolutionary pressure and expansion in all three primate species, while some are under expansion uniquely in human, chimpanzee, or macaque. For example, cholesterol pathways are conserved in all three species (and other primate species). In all three species, immune response genes are under positive selection, and genes of T cell-mediated immunity, signal transduction, cell adhesion, and membrane proteins generally. Genes for keratin, which produce hair shafts, were rapidly evolving in all three species, possibly because of climate change or mate selection. The X chromosome has three times more rearrangements than other chromosomes. The macaque gained 1,358 genes by duplication. Triangulation of human, chimpanzee, and macaque sequences showed expansion of gene families in each species. The ''PKFP'' gene, important in sugar (fructose) metabolism, is expanded in macaques, possibly because of their high-
fruit In botany, a fruit is the seed-bearing structure in flowering plants that is formed from the ovary after flowering. Fruits are the means by which flowering plants (also known as angiosperms) disseminate their seeds. Edible fruits in particu ...
diet. So are genes for the olfactory receptor, cytochrome P450 (which degrades toxins), and CCL3L1-CCL4 (associated in humans with HIV susceptibility). Immune genes are expanded in macaques, relative to all four
great ape The Hominidae (), whose members are known as the great apes or hominids (), are a taxonomic family of primates that includes eight extant species in four genera: '' Pongo'' (the Bornean, Sumatran and Tapanuli orangutan); ''Gorilla'' (the ...
species. The macaque genome has 33 major histocompatibility genes, three times those of human. This has clinical significance because the macaque is used as an experimental model of the human immune system. In humans, the preferentially expressed antigen of melanoma (PRAME) gene family is expanded. It is actively expressed in cancers, but normally is testis-specific, possibly involved in spermatogenesis. The PRAME family has 26 members on human chromosome 1. In the macaque, it has eight, and has been very simple and stable for millions of years. The PRAME family arose in translocations in the common mouse-primate ancestor 85 million years ago, and is expanded on mouse chromosome 4. DNA microarrays are used in macaque research. For example,
Michael Katze Michael Katze was an American microbiologist. For over 35 years, he has researched host-virus interactions, incorporating systems biology approaches into infectious disease research.Katze Lab: People'Katze Biosketch' He was an international leader ...
of University of Washington, Seattle, infected macaques with 1918 and modern influenzas. The DNA microarray showed the macaque genomic response to human influenza on a cellular level in each tissue. Both viruses stimulated innate immune system inflammation, but the 1918 flu stimulated stronger and more persistent inflammation, causing extensive tissue damage, and it did not stimulate the interferon-1 pathway. The DNA response showed a transition from innate to adaptive immune response over seven days. The full sequence and annotation of the macaque genome is available on the
Ensembl Ensembl genome database project is a scientific project at the European Bioinformatics Institute, which provides a centralized resource for geneticists, molecular biologists and other researchers studying the genomes of our own species and other v ...
genome browser.


Conservation Status

The rhesus macaque is listed as
Least Concern A least-concern species is a species that has been categorized by the International Union for Conservation of Nature (IUCN) as evaluated as not being a focus of species conservation because the specific species is still plentiful in the wild. T ...
on the
IUCN Red List The International Union for Conservation of Nature (IUCN) Red List of Threatened Species, also known as the IUCN Red List or Red Data Book, founded in 1964, is the world's most comprehensive inventory of the global conservation status of biol ...
and estimated to exist in large numbers; it is tolerant of a broad range of
habitat In ecology, the term habitat summarises the array of resources, physical and biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species habitat can be seen as the physical ...
s, including urban environments. Rhesus macaques have the largest natural range of any non-human primate which contributes to the conservation status of "least concern". The Thai population is locally threatened. In addition to habitat destruction and agricultural encroachment, pet releases of the different species into existing troops are diluting the gene pool and putting its genetic integrity at risk. Despite the wealth of information on their ecology and behaviour, little attention has been paid to their demography or population status, which can pose a risk for future rhesus macaque population. Rhesus macaques increased population stress on other species, having extended their distributional limits by approximately 3,500 km2 in Southeastern India. The increased area of rhesus macaques has been caused by human intervention tactics whereby village translocation occurs from urban conflict ridden areas. This influx has led to the widespread establishment of the rhesus macaque, accompanied by the disappearance of the bonnet macaque in these areas.


Trivia

An archaic name for the rhesus macaque is bruh. This, however, is not a common name in contemporary times.


See also

*
Girneys Girneys are soft vocalizations used by species of Old World monkeys to ease affiliative social interactions between unrelated members of the same species. The vocalizations are most commonly used by adult females around birthing season; the fem ...


References


External links


ARKive: images and movies of the rhesus macaque ''Macaca mulatta''

Brain maps and brain atlases of rhesus macaque

Primate Info Net: ''Macaca mulatta'' Factsheet


* ttp://www.rhesusgenome.org ''Macaca mulatta'' Genome
Rhesus Play
Film analysis of agonistic play by Donald Symons (UCSB) on DVD * View th
Macaque genome
in
Ensembl Ensembl genome database project is a scientific project at the European Bioinformatics Institute, which provides a centralized resource for geneticists, molecular biologists and other researchers studying the genomes of our own species and other v ...
. * {{Authority control Animal models Fauna of Yunnan
rhesus macaque The rhesus macaque (''Macaca mulatta''), colloquially rhesus monkey, is a species of Old World monkey. There are between six and nine recognised subspecies that are split between two groups, the Chinese-derived and the Indian-derived. Generally ...
rhesus macaque The rhesus macaque (''Macaca mulatta''), colloquially rhesus monkey, is a species of Old World monkey. There are between six and nine recognised subspecies that are split between two groups, the Chinese-derived and the Indian-derived. Generally ...
Mammals of Afghanistan Mammals of Bangladesh Mammals of Bhutan Mammals of China Mammals of Hong Kong Mammals of India Mammals of Laos Mammals of Myanmar Mammals of Nepal Mammals of Pakistan Mammals of Thailand Mammals of Vietnam Monkeys in India Primates of Asia Space-flown life Taxa named by Eberhard August Wilhelm von Zimmermann