Rhesus Macaque
Overview
The Rhesus macaque also called the Rhesus monkey, is one of the best-known species of Old World monkey
Old World monkey
The Old World monkeys or Cercopithecidae are a group of primates, falling in the superfamily Cercopithecoidea in the clade Catarrhini. The Old World monkeys are native to Africa and Asia today, inhabiting a range of environments from tropical rain forest to savanna, shrubland and mountainous...

s. It is listed as Least Concern
Least Concern
Least Concern is an IUCN category assigned to extant taxon or lower taxa which have been evaluated but do not qualify for any other category. As such they do not qualify as threatened, Near Threatened, or Conservation Dependent...

 in the IUCN Red List of Threatened Species in view of its wide distribution, presumed large population, and its tolerance of a broad range of habitats. Native to South
South Asia
South Asia, also known as Southern Asia, is the southern region of the Asian continent, which comprises the sub-Himalayan countries and, for some authorities , also includes the adjoining countries to the west and the east...

, Central
Central Asia
Central Asia is a core region of the Asian continent from the Caspian Sea in the west, China in the east, Afghanistan in the south, and Russia in the north...

 and Southeast Asia
Southeast Asia
Southeast Asia, South-East Asia, South East Asia or Southeastern Asia is a subregion of Asia, consisting of the countries that are geographically south of China, east of India, west of New Guinea and north of Australia. The region lies on the intersection of geological plates, with heavy seismic...

, troops of Macaca mulatta inhabit a great variety of habitats from grasslands to arid and forested areas, but also close to human settlements.
The Rhesus macaque is brown or grey in color and has a pink face, which is bereft of fur.
Encyclopedia
The Rhesus macaque also called the Rhesus monkey, is one of the best-known species of Old World monkey
Old World monkey
The Old World monkeys or Cercopithecidae are a group of primates, falling in the superfamily Cercopithecoidea in the clade Catarrhini. The Old World monkeys are native to Africa and Asia today, inhabiting a range of environments from tropical rain forest to savanna, shrubland and mountainous...

s. It is listed as Least Concern
Least Concern
Least Concern is an IUCN category assigned to extant taxon or lower taxa which have been evaluated but do not qualify for any other category. As such they do not qualify as threatened, Near Threatened, or Conservation Dependent...

 in the IUCN Red List of Threatened Species in view of its wide distribution, presumed large population, and its tolerance of a broad range of habitats. Native to South
South Asia
South Asia, also known as Southern Asia, is the southern region of the Asian continent, which comprises the sub-Himalayan countries and, for some authorities , also includes the adjoining countries to the west and the east...

, Central
Central Asia
Central Asia is a core region of the Asian continent from the Caspian Sea in the west, China in the east, Afghanistan in the south, and Russia in the north...

 and Southeast Asia
Southeast Asia
Southeast Asia, South-East Asia, South East Asia or Southeastern Asia is a subregion of Asia, consisting of the countries that are geographically south of China, east of India, west of New Guinea and north of Australia. The region lies on the intersection of geological plates, with heavy seismic...

, troops of Macaca mulatta inhabit a great variety of habitats from grasslands to arid and forested areas, but also close to human settlements.

Characteristics

The Rhesus macaque is brown or grey in color and has a pink face, which is bereft of fur. Its tail is of medium length and averages between 20.7 and 22.9 cm (8.1 and 9 in). Adult males measure approximately 53 cm (20.9 in) on average and weigh about 7.7 kg (17 lb). Females are smaller, averaging 47 cm (18.5 in) in length and 5.3 kg (11.7 lb) in weight. Rhesus macaques have on average 50 vertebrae. Their intermembral index (ratio of arms to legs) is 89%. They have dorsal scapula
Scapula
In anatomy, the scapula , omo, or shoulder blade, is the bone that connects the humerus with the clavicle ....

 and a wide rib cage.

The Rhesus macaque has 32 teeth with a dental formula of 2.1.2.3/2.1.2.3 and bilophodont molars. The upper molars have four cusp
Cusp
Cusp may refer to:*Beach cusps, a pointed and regular arc pattern of the shoreline at the beach*Behavioral cusp an important behavior change with far reaching consequences*Cusp catastrophe...

s: paracone, metacone
Metacone
thumb|500px|left|Right upper molar showing the four main upper molars cusps.The metacone is a cusp on the molars of the upper dentition in hominids. It is found at the buccal distal area of the tooth...

, protocone and hypocone. The lower molars also have four cusps: metaconid, protoconid, hypoconid and entoconid.

Distribution and habitat

Rhesus macaques are native to northern India
India
India , officially the Republic of India , is a country in South Asia. It is the seventh-largest country by geographical area, the second-most populous country with over 1.2 billion people, and the most populous democracy in the world...

, Bangladesh
Bangladesh
Bangladesh , officially the People's Republic of Bangladesh is a sovereign state located in South Asia. It is bordered by India on all sides except for a small border with Burma to the far southeast and by the Bay of Bengal to the south...

, Pakistan
Pakistan
Pakistan , officially the Islamic Republic of Pakistan is a sovereign state in South Asia. It has a coastline along the Arabian Sea and the Gulf of Oman in the south and is bordered by Afghanistan and Iran in the west, India in the east and China in the far northeast. In the north, Tajikistan...

, Nepal
Nepal
Nepal , officially the Federal Democratic Republic of Nepal, is a landlocked sovereign state located in South Asia. It is located in the Himalayas and bordered to the north by the People's Republic of China, and to the south, east, and west by the Republic of India...

, Burma, Thailand
Thailand
Thailand , officially the Kingdom of Thailand , formerly known as Siam , is a country located at the centre of the Indochina peninsula and Southeast Asia. It is bordered to the north by Burma and Laos, to the east by Laos and Cambodia, to the south by the Gulf of Thailand and Malaysia, and to the...

, Afghanistan
Afghanistan
Afghanistan , officially the Islamic Republic of Afghanistan, is a landlocked country located in the centre of Asia, forming South Asia, Central Asia and the Middle East. With a population of about 29 million, it has an area of , making it the 42nd most populous and 41st largest nation in the world...

, Vietnam
Vietnam
Vietnam – sometimes spelled Viet Nam , officially the Socialist Republic of Vietnam – is the easternmost country on the Indochina Peninsula in Southeast Asia. It is bordered by China to the north, Laos to the northwest, Cambodia to the southwest, and the South China Sea –...

, southern China
China
Chinese civilization may refer to:* China for more general discussion of the country.* Chinese culture* Greater China, the transnational community of ethnic Chinese.* History of China* Sinosphere, the area historically affected by Chinese culture...

, and some neighboring areas. The Rhesus monkey has the widest geographic ranges of any nonhuman 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 2500 m (8,202.1 ft) in elevation. They are regular swimmers. Babies as young as a few days old can swim, and adults are known to swim over a half mile between islands, but are often found drowned in small groups where their drinking waters lie. Rhesus macaques are noted for their tendency to move from rural to urban areas, coming to rely on handouts or refuse from humans.

The southern and the northern distributional limits for rhesus and bonnet macaque
Bonnet Macaque
The bonnet macaque is a macaque endemic to southern India. Its distribution is limited by the Indian Ocean on three sides and the Godavari and Tapti Rivers along with a related competing species of rhesus macaque in the north....

s, respectively, currently run parallel to each other in the western part of the country, are separated by a large gap in central India, 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.

Distribution of subspecies and populations

The name rhesus is reminiscent of the Greek mythological king Rhêsos
Rhesus of Thrace
Rhesus or Rhêsos was a Thracian king who fought on the side of Trojans in Iliad, Book X, where Diomedes and Odysseus stole his team of fine horses during a night raid on the Trojan camp. Homer gives his father as Eioneus— a name otherwise given to the father of Dia, whom Ixion threw into the...

. However, the French naturalist Jean-Baptiste Audebert
Jean-Baptiste Audebert
Jean Baptiste Audebert was a French artist and naturalist.Audebert was born at Rochefort. He studied painting and drawing at Paris, and gained considerable reputation as a miniature-painter. Employed in preparing plates for the Histoire des cloportes of Guillaume-Antoine Olivier, he acquired a...

 who applied the name to the species, stated that it had no meaning.

According to Zimmermann’s first description of 1780, the Rhesus macaque is distributed in eastern Afghanistan
Afghanistan
Afghanistan , officially the Islamic Republic of Afghanistan, is a landlocked country located in the centre of Asia, forming South Asia, Central Asia and the Middle East. With a population of about 29 million, it has an area of , making it the 42nd most populous and 41st largest nation in the world...

, Bangladesh
Bangladesh
Bangladesh , officially the People's Republic of Bangladesh is a sovereign state located in South Asia. It is bordered by India on all sides except for a small border with Burma to the far southeast and by the Bay of Bengal to the south...

, Bhutan
Bhutan
Bhutan , officially the Kingdom of Bhutan, is a landlocked state in South Asia, located at the eastern end of the Himalayas and bordered to the south, east and west by the Republic of India and to the north by the People's Republic of China...

, as far east as the Brahmaputra Valley in peninsular India
India
India , officially the Republic of India , is a country in South Asia. It is the seventh-largest country by geographical area, the second-most populous country with over 1.2 billion people, and the most populous democracy in the world...

, Nepal
Nepal
Nepal , officially the Federal Democratic Republic of Nepal, is a landlocked sovereign state located in South Asia. It is located in the Himalayas and bordered to the north by the People's Republic of China, and to the south, east, and west by the Republic of India...

 and northern Pakistan
Pakistan
Pakistan , officially the Islamic Republic of Pakistan is a sovereign state in South Asia. It has a coastline along the Arabian Sea and the Gulf of Oman in the south and is bordered by Afghanistan and Iran in the west, India in the east and China in the far northeast. In the north, Tajikistan...

. Today, this is known as the Indian Rhesus macaque Macaca mulatta mulatta, which includes the morphologically similar Macacus rhesus villosus described by True
Frederick W. True
Frederick William True was an American biologist, the first head curator of biology at the United States National Museum, now part of the Smithsonian Institution....

 in 1894 from Kashmir
Kashmir
Kashmir is the northwestern region of the Indian subcontinent. Until the mid-19th century, the term Kashmir geographically denoted only the valley between the Great Himalayas and the Pir Panjal mountain range...

 and Macaca mulatta mcmahoni described by Pocock
Reginald Innes Pocock
Reginald Innes Pocock F.R.S. was a British zoologist.Pocock was born in Clifton, Bristol, the fourth son of Rev. Nicholas Pocock and Edith Prichard. He began showing interest in natural history at St. Edward's School, Oxford. He received tutoring in zoology from Sir Edward Poulton, and was allowed...

 in 1932 from Kootai, Pakistan
Pakistan
Pakistan , officially the Islamic Republic of Pakistan is a sovereign state in South Asia. It has a coastline along the Arabian Sea and the Gulf of Oman in the south and is bordered by Afghanistan and Iran in the west, India in the east and China in the far northeast. In the north, Tajikistan...

. Several Chinese
China
Chinese civilization may refer to:* China for more general discussion of the country.* Chinese culture* Greater China, the transnational community of ethnic Chinese.* History of China* Sinosphere, the area historically affected by Chinese culture...

 subspecies
Subspecies
Subspecies in biological classification, is either a taxonomic rank subordinate to species, ora taxonomic unit in that rank . A subspecies cannot be recognized in isolation: a species will either be recognized as having no subspecies at all or two or more, never just one...

 of Rhesus macaques have been 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 in:
  • Macaca mulatta mulatta – is found in western and central China, in the south of Yunnan
    Yunnan
    Yunnan is a province of the People's Republic of China, located in the far southwest of the country spanning approximately and with a population of 45.7 million . The capital of the province is Kunming. The province borders Burma, Laos, and Vietnam.Yunnan is situated in a mountainous area, with...

     and southwest of Guangxi
    Guangxi
    Guangxi, formerly romanized Kwangsi, is a province of southern China along its border with Vietnam. In 1958, it became the Guangxi Zhuang Autonomous Region of the People's Republic of China, a region with special privileges created specifically for the Zhuang people.Guangxi's location, in...

    ;
  • Macaca mulatta lasiota (Gray
    John Edward Gray
    John Edward Gray, FRS was a British zoologist. He was the elder brother of George Robert Gray and son of the pharmacologist and botanist Samuel Frederick Gray ....

    , 1868) – the West Chinese Rhesus macaque is distributed in the west of Sichuan
    Sichuan
    ' , known formerly in the West by its postal map spellings of Szechwan or Szechuan is a province in Southwest China with its capital in Chengdu...

    , northwest of Yunnan
    Yunnan
    Yunnan is a province of the People's Republic of China, located in the far southwest of the country spanning approximately and with a population of 45.7 million . The capital of the province is Kunming. The province borders Burma, Laos, and Vietnam.Yunnan is situated in a mountainous area, with...

    , southeast of Qinghai
    Qinghai
    Qinghai ; Oirat Mongolian: ; ; Salar:) is a province of the People's Republic of China, named after Qinghai Lake...

    ; possibly synonymous with Macaca mulatta sanctijohannis (Swinhoe
    Robert Swinhoe
    Robert Swinhoe FRS , was an English naturalist who worked as a Consul in Formosa. He discovered many Southeast Asian birds and several, such as Swinhoe's Pheasant, are named after him.-Biography:...

    , 1867), if not with Macaca mulatta mulatta.
  • Macaca mulatta tcheliensis (Milne-Edwards
    Henri Milne-Edwards
    Henri Milne-Edwards was an eminent French zoologist.Henri Milne-Edwards was the 27th child of William Edwards, an English planter and militia colonel in Jamaica and Elisabeth Vaux, a French. He was born in Bruges, Belgium, where his parents had retired. At that time, Bruges was a part of the...

    , 1870) – the North Chinese Rhesus macaque lives in the north of Henan
    Henan
    Henan , is a province of the People's Republic of China, located in the central part of the country. Its one-character abbreviation is "豫" , named after Yuzhou , a Han Dynasty state that included parts of Henan...

    , south of Shanxi
    Shanxi
    ' is a province in Northern China. Its one-character abbreviation is "晋" , after the state of Jin that existed here during the Spring and Autumn Period....

     and near Beijing
    Beijing
    Beijing , also known as Peking , is the capital of the People's Republic of China and one of the most populous cities in the world, with a population of 19,612,368 as of 2010. The city is the country's political, cultural, and educational center, and home to the headquarters for most of China's...

    . Some consider it as the most endangered subspecies. Others consider it possibly synonymous with M. m. sanctijohannis, if not with M. m. mulatta.
  • Macaca mulatta vestita (Milne-Edwards
    Henri Milne-Edwards
    Henri Milne-Edwards was an eminent French zoologist.Henri Milne-Edwards was the 27th child of William Edwards, an English planter and militia colonel in Jamaica and Elisabeth Vaux, a French. He was born in Bruges, Belgium, where his parents had retired. At that time, Bruges was a part of the...

    , 1892) – the Tibetan Rhesus macaque lives in the southeast of Tibet
    Tibet
    Tibet is a plateau region in Asia, north-east of the Himalayas. It is the traditional homeland of the Tibetan people as well as some other ethnic groups such as Monpas, Qiang, and Lhobas, and is now also inhabited by considerable numbers of Han and Hui people...

    , northwest of Yunnan (Deqing), and perhaps including Yushu
    Yushu
    Yushu may refer to the following locations in China:*Yushu Tibetan Autonomous Prefecture , in Qinghai*Yushu, Jilin , city in Jilin*Yushu County , in Qinghai*Yushu Town, the name of a number of towns...

    ; possibly synonymous with M. m. sanctijohannis, if not with M. m. mulatta.
  • Macaca mulatta littoralis (Elliot
    Daniel Giraud Elliot
    Daniel Giraud Elliot was an American zoologist.Elliot was one of the founders of the American Museum of Natural History in New York and the American Ornithologists' Union. He was also curator of zoology at the Field Museum in Chicago.Elliot used his wealth to publish a series of sumptuous...

    , 1909) – the South Chinese Rhesus macaque lives in Fujian
    Fujian
    ' , formerly romanised as Fukien or Huguing or Foukien, is a province on the southeast coast of mainland China. Fujian is bordered by Zhejiang to the north, Jiangxi to the west, and Guangdong to the south. Taiwan lies to the east, across the Taiwan Strait...

    , Zhejiang
    Zhejiang
    Zhejiang is an eastern coastal province of the People's Republic of China. The word Zhejiang was the old name of the Qiantang River, which passes through Hangzhou, the provincial capital...

    , Anhui
    Anhui
    Anhui is a province in the People's Republic of China. Located in eastern China across the basins of the Yangtze River and the Huai River, it borders Jiangsu to the east, Zhejiang to the southeast, Jiangxi to the south, Hubei to the southwest, Henan to the northwest, and Shandong for a tiny...

    , Jiangxi
    Jiangxi
    ' is a southern province in the People's Republic of China. Spanning from the banks of the Yangtze River in the north into hillier areas in the south, it shares a border with Anhui to the north, Zhejiang to the northeast, Fujian to the east, Guangdong to the south, Hunan to the west, and Hubei to...

    , Hunan
    Hunan
    ' is a province of South-Central China, located to the south of the middle reaches of the Yangtze River and south of Lake Dongting...

    , Hubei
    Hubei
    ' Hupeh) is a province in Central China. The name of the province means "north of the lake", referring to its position north of Lake Dongting...

    , Guizhou
    Guizhou
    ' is a province of the People's Republic of China located in the southwestern part of the country. Its provincial capital city is Guiyang.- History :...

    , northwest of Guangdong
    Guangdong
    Guangdong is a province on the South China Sea coast of the People's Republic of China. The province was previously often written with the alternative English name Kwangtung Province...

    , north of Guangxi
    Guangxi
    Guangxi, formerly romanized Kwangsi, is a province of southern China along its border with Vietnam. In 1958, it became the Guangxi Zhuang Autonomous Region of the People's Republic of China, a region with special privileges created specifically for the Zhuang people.Guangxi's location, in...

    , northeast of Yunnan
    Yunnan
    Yunnan is a province of the People's Republic of China, located in the far southwest of the country spanning approximately and with a population of 45.7 million . The capital of the province is Kunming. The province borders Burma, Laos, and Vietnam.Yunnan is situated in a mountainous area, with...

    , east of Sichuan
    Sichuan
    ' , known formerly in the West by its postal map spellings of Szechwan or Szechuan is a province in Southwest China with its capital in Chengdu...

     and south of Shaanxi
    Shaanxi
    ' is a province in the central part of Mainland China, and it includes portions of the Loess Plateau on the middle reaches of the Yellow River in addition to the Qinling Mountains across the southern part of this province...

    ; possibly synonymous with M. m. sanctijohannis, if not with M. m. mulatta.
  • Macaca mulatta brevicaudus, also referred to as Pithecus brevicaudus (Elliot
    Daniel Giraud Elliot
    Daniel Giraud Elliot was an American zoologist.Elliot was one of the founders of the American Museum of Natural History in New York and the American Ornithologists' Union. He was also curator of zoology at the Field Museum in Chicago.Elliot used his wealth to publish a series of sumptuous...

    , 1913) – lives in the Hainan Island and Wanshan Islands in Guangdong
    Guangdong
    Guangdong is a province on the South China Sea coast of the People's Republic of China. The province was previously often written with the alternative English name Kwangtung Province...

    , and the islands near Hong Kong
    Hong Kong
    Hong Kong is one of two Special Administrative Regions of the People's Republic of China , the other being Macau. A city-state situated on China's south coast and enclosed by the Pearl River Delta and South China Sea, it is renowned for its expansive skyline and deep natural harbour...

    ; may be synonymous with M. m. mulatta.
  • Macaca mulatta siamica (Kloss
    Cecil Boden Kloss
    Cecil Boden Kloss was an English zoologist. He was an expert on the mammals and birds of Southeast Asia.In the early 1900s Kloss accompanied the American naturalist William Louis Abbott in exploring the Andaman and Nicobar islands. From 1908 he worked under Herbert Christopher Robinson at the...

    , 1917) – the Indochinese Rhesus macaque is distributed in Myanmar
    Myanmar
    Burma , officially the Republic of the Union of Myanmar , is a country in Southeast Asia. Burma is bordered by China on the northeast, Laos on the east, Thailand on the southeast, Bangladesh on the west, India on the northwest, the Bay of Bengal to the southwest, and the Andaman Sea on the south....

    , in the north of Thailand
    Thailand
    Thailand , officially the Kingdom of Thailand , formerly known as Siam , is a country located at the centre of the Indochina peninsula and Southeast Asia. It is bordered to the north by Burma and Laos, to the east by Laos and Cambodia, to the south by the Gulf of Thailand and Malaysia, and to the...

     and Vietnam
    Vietnam
    Vietnam – sometimes spelled Viet Nam , officially the Socialist Republic of Vietnam – is the easternmost country on the Indochina Peninsula in Southeast Asia. It is bordered by China to the north, Laos to the northwest, Cambodia to the southwest, and the South China Sea –...

    , in Laos
    Laos
    Laos Lao: ສາທາລະນະລັດ ປະຊາທິປະໄຕ ປະຊາຊົນລາວ Sathalanalat Paxathipatai Paxaxon Lao, officially the Lao People's Democratic Republic, is a landlocked country in Southeast Asia, bordered by Burma and China to the northwest, Vietnam to the east, Cambodia to the south and Thailand to the west...

     and in the Chinese provinces of Anhui
    Anhui
    Anhui is a province in the People's Republic of China. Located in eastern China across the basins of the Yangtze River and the Huai River, it borders Jiangsu to the east, Zhejiang to the southeast, Jiangxi to the south, Hubei to the southwest, Henan to the northwest, and Shandong for a tiny...

    , northwest Guangxi
    Guangxi
    Guangxi, formerly romanized Kwangsi, is a province of southern China along its border with Vietnam. In 1958, it became the Guangxi Zhuang Autonomous Region of the People's Republic of China, a region with special privileges created specifically for the Zhuang people.Guangxi's location, in...

    , Guizhou
    Guizhou
    ' is a province of the People's Republic of China located in the southwestern part of the country. Its provincial capital city is Guiyang.- History :...

    , Hubei
    Hubei
    ' Hupeh) is a province in Central China. The name of the province means "north of the lake", referring to its position north of Lake Dongting...

    , Hunan
    Hunan
    ' is a province of South-Central China, located to the south of the middle reaches of the Yangtze River and south of Lake Dongting...

    , central and eastern Sichuan
    Sichuan
    ' , known formerly in the West by its postal map spellings of Szechwan or Szechuan is a province in Southwest China with its capital in Chengdu...

    , western and south-central Yunnan
    Yunnan
    Yunnan is a province of the People's Republic of China, located in the far southwest of the country spanning approximately and with a population of 45.7 million . The capital of the province is Kunming. The province borders Burma, Laos, and Vietnam.Yunnan is situated in a mountainous area, with...

    ; possibly synonymous with M. m. sanctijohannis, if not with M. m. mulatta.

Feral colonies in the United States

Around the spring of 1938, a colony of Rhesus macaques were released in the Silver River State Park
Silver River State Park
Silver River State Park is a nearly 5,000 acre Florida State Park east of Ocala. Adjacent to Silver Springs and the Ocklawaha River, the park is a mile south of State Road 40 on State Road 35 .-Ecology:...

 in Florida
Florida
Florida is a state in the southeastern United States, located on the nation's Atlantic and Gulf coasts. It is bordered to the west by the Gulf of Mexico, to the north by Alabama and Georgia and to the east by the Atlantic Ocean. With a population of 18,801,310 as measured by the 2010 census, it...

 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
Tarzan is a fictional character, an archetypal feral child raised in the African jungles by the Mangani "great apes"; he later experiences civilization only to largely reject it and return to the wild as a heroic adventurer...

 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!
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".-Plot:...

contains no Rhesus macaques. In addition, various colonies of Rhesus and other monkey species are speculated to be the result of zoos and wildlife parks destroyed in hurricanes, most notably Hurricane Andrew
Hurricane Andrew
Hurricane Andrew was the third Category 5 hurricane to make landfall in the United States, after the Labor Day Hurricane of 1935 and Hurricane Camille in 1969. Andrew was the first named storm and only major hurricane of the otherwise inactive 1992 Atlantic hurricane season...

.

There is also a notable colony of Rhesus macaques on Morgan Island, one of the Sea Islands
Sea Islands
The Sea Islands are a chain of tidal and barrier islands on the Atlantic Ocean coast of the United States. They number over 100, and are located between the mouths of the Santee and St. Johns Rivers along the coast of the U.S...

 in the South Carolina Lowcountry
South Carolina Lowcountry
The Lowcountry is a geographic and cultural region located along South Carolina's coast. The region includes the South Carolina Sea Islands...

. They were imported in the 1970s for use in local labs and are by all accounts thriving.

Ecology and behavior

Rhesus macaques are diurnal animal
Diurnal animal
Diurnality is a plant or animal behavior characterized by activity during the day and sleeping at night.-In animals:Animals that are not diurnal might be nocturnal or crepuscular . Many animal species are diurnal, including many mammals, insects, reptiles and birds...

s, and both arboreal and terrestrial
Terrestrial animal
Terrestrial animals are animals that live predominantly or entirely on land , as compared with aquatic animals, which live predominantly or entirely in the water , or amphibians, which rely on a combination of aquatic and terrestrial habitats...

. They are quadrupedal and, when on the ground, they walk digitigrade and plantigrade. They are mostly herbivorous feeding on mainly fruit
Fruit
In broad terms, a fruit is a structure of a plant that contains its seeds.The term has different meanings dependent on context. In non-technical usage, such as food preparation, fruit normally means the fleshy seed-associated structures of certain plants that are sweet and edible in the raw state,...

, but also eating seed
Seed
A seed is a small embryonic plant enclosed in a covering called the seed coat, usually with some stored food. It is the product of the ripened ovule of gymnosperm and angiosperm plants which occurs after fertilization and some growth within the mother plant...

s, root
Root
In vascular plants, the root is the organ of a plant that typically lies below the surface of the soil. This is not always the case, however, since a root can also be aerial or aerating . Furthermore, a stem normally occurring below ground is not exceptional either...

s, buds, bark
Bark
Bark is the outermost layers of stems and roots of woody plants. Plants with bark include trees, woody vines and shrubs. Bark refers to all the tissues outside of the vascular cambium and is a nontechnical term. It overlays the wood and consists of the inner bark and the outer bark. The inner...

, and cereal
Cereal
Cereals are grasses cultivated for the edible components of their grain , composed of the endosperm, germ, and bran...

s. It has been estimated that they consume around 99 different plant species in 46 families. During the monsoon
Monsoon
Monsoon is traditionally defined as 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 the asymmetric heating of land and sea...

 season, they get much of their water from ripe and succulent fruit. Macaque living far from water sources lick dewdrops from leaves and drink rain water accumulated in tree hollows. They have also been observed eating termite
Termite
Termites are a group of eusocial insects that, until recently, were classified at the taxonomic rank of order Isoptera , but are now accepted as the epifamily Termitoidae, of the cockroach order Blattodea...

s, grasshopper
Grasshopper
The grasshopper is an insect of the suborder Caelifera in the order Orthoptera. To distinguish it from bush crickets or katydids, it is sometimes referred to as the short-horned grasshopper...

s, ant
Ant
Ants are social insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from wasp-like ancestors in the mid-Cretaceous period between 110 and 130 million years ago and diversified after the rise of flowering plants. More than...

s and beetle
Beetle
Coleoptera is an order of insects commonly called beetles. The word "coleoptera" is from the Greek , koleos, "sheath"; and , pteron, "wing", thus "sheathed wing". Coleoptera contains more species than any other order, constituting almost 25% of all known life-forms...

s. When food is abundant, they are distributed in patches and forage throughout the day in their home range. 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
Comparative psychology
Comparative psychology generally refers to the scientific study of the behavior and mental processes of non-human animals. However, scientists from different disciplines do not always agree on this definition...

 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, an important type of self-awareness.

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. Females have highly stable matrilineal 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 that exist 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 their early years, rebelling against their mother is discouraged. Juvenile male macaques also exist in matrilineal lines but once they reach 4–5 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 Sub-group contains the 2-3 oldest and most dominant males who are co-dominant, along with females, their infants and juveniles. This sub-group occupies the center of the group and determines the movements, foraging and other routines. The females of this sub-group are also the most dominant of the entire group. The farther to the periphery a sub-group is, the less dominant it is. Sub-groups on the periphery of the central group are run by one dominant male who ranks lower than the central males, and who maintain order in the group and communicate messages between the central and peripheral males. A sub-group of subordinate, often sub-adult males occupy the very edge of the groups and have the responsibility of communicating with other macaque groups and make alarm calls.

Communication

Rhesus macaques interact using a variety of facial expressive, vocalizations and 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 rank with the lower rank one giving the expression to its superior. A less dominant individual will also make 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 will threaten another individual standing quadrupedally making a silent "open mouth stare" accompanied by the tail sticking straight. During movements, macaques will make "coos" and "grunts". These are also made during affiliative interactions and approaches before grooming. When they find rare food of high-quality, macaque will emit "Warbles," "harmonic arches," or "chirps." When in threatening situations, macaques will emit a single loud, high-pitched sounds called a "shrill bark". "Screeches," "screams," "squeaks," "pant-threats," "growls," and "barks" are when during aggressive interactions. Infants make "geckers" during weaning conflicts.

Reproduction

Adult male macaques try to maximize their reproductive success by entering into consort pairs with females, both in and outside the breeding period. Females prefer to mate with males that will increase the survival of their young. Thus a consort male provides resources for his female and protects her from predators. Larger, more dominant males are more likely to provide for the females. The breeding period can last up to 11 days, and a female usually mates with four males during that time. Male rhesus macaques have not been observed to fight for access to sexually receptive females, although they suffer more wounds during the mating season. Female macaques first breed when they are four years old, and reach menopause
Menopause
Menopause is a term used to describe the permanent cessation of the primary functions of the human ovaries: the ripening and release of ova and the release of hormones that cause both the creation of the uterine lining and the subsequent shedding of the uterine lining...

 at around 25 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.

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. This is because the mothers may pass the parenting responsibilities to her 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 the preceding birth season. Infants farther from the center of the groups are more vulnerable to infanticide from outside groups. Some mothers abuse their infants which is believed to be the result of controlling parenting styles.

In science

The Rhesus macaque is well known to science. Due to its relatively easy upkeep in captivity, its wide availability and its 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 Rhesus factor, one of the elements of a person's blood group
Blood type
A blood type is a classification of blood based on the presence or absence of inherited antigenic substances on the surface of red blood cells . These antigens may be proteins, carbohydrates, glycoproteins, or glycolipids, depending on the blood group system...

, by the discoverers of the factor, Karl Landsteiner
Karl Landsteiner
Karl Landsteiner , was an Austrian-born American biologist and physician of Jewish origin. He is noted for having first distinguished the main blood groups in 1900, having developed the modern system of classification of blood groups from his identification of the presence of agglutinins in the...

 and Alexander Wiener. The Rhesus macaque was also used in the well-known experiments on maternal deprivation carried out in the 1950s by controversial
Bioethics
Bioethics is the study of controversial ethics brought about by advances in biology and medicine. Bioethicists are concerned with the ethical questions that arise in the relationships among life sciences, biotechnology, medicine, politics, law, and philosophy....

 comparative psychologist
Comparative psychology
Comparative psychology generally refers to the scientific study of the behavior and mental processes of non-human animals. However, scientists from different disciplines do not always agree on this definition...

 Harry Harlow
Harry Harlow
Harry Frederick Harlow was an American psychologist best known for his maternal-separation and social isolation experiments on rhesus monkeys, which demonstrated the importance of care-giving and companionship in social and cognitive development...

. Other medical breakthroughs facilitated by the use of the Rhesus macaque include:
  • development of the rabies
    Rabies
    Rabies is a viral disease that causes acute encephalitis in warm-blooded animals. It is zoonotic , most commonly by a bite from an infected animal. For a human, rabies is almost invariably fatal if post-exposure prophylaxis is not administered prior to the onset of severe symptoms...

    , smallpox
    Smallpox
    Smallpox was an infectious disease unique to humans, caused by either of two virus variants, Variola major and Variola minor. The disease is also known by the Latin names Variola or Variola vera, which is a derivative of the Latin varius, meaning "spotted", or varus, meaning "pimple"...

    , and polio vaccines
  • creation of drugs to manage HIV/AIDS
  • understanding of the female reproductive cycle and development of the embryo and the propagation of embryonic stem cells.


The U.S. Army, the U.S. Air Force, and NASA
NASA
The National Aeronautics and Space Administration is the agency of the United States government that is responsible for the nation's civilian space program and for aeronautics and aerospace research...

 launched Rhesus macaques into outer space during the 1950s and 60s, and the Soviet/Russian space program launched them into space as recently as 1997 on the Bion missions. One of these primates ("Able") who was launched on a suborbital spaceflight
Monkeys in space
Before humans went into space, several animals were launched into space, including numerous monkeys, so that scientists could investigate the biological effects of space travel. The United States launched flights containing primate cargo primarily between 1948-1961 with one flight in 1969 and one...

 in 1959 was one of the two first living beings (along with "Miss Baker" on the same mission) to travel in space and return alive.

In January 2000, the Rhesus macaque became the first cloned
Cloning
Cloning in biology is the process of producing similar populations of genetically identical individuals that occurs in nature when organisms such as bacteria, insects or plants reproduce asexually. Cloning in biotechnology refers to processes used to create copies of DNA fragments , cells , or...

 primate
Primate
A primate is a mammal of the order Primates , which contains prosimians and simians. Primates arose from ancestors that lived in the trees of tropical forests; many primate characteristics represent adaptations to life in this challenging three-dimensional environment...

 with the birth of Tetra. January 2001 saw the birth of ANDi
ANDI
Andi or ANDI may refer to:*Andi, Yinan County , town in Yinan County, Shandong, China*Andi, Estonia, village in Vihula Parish, Lääne-Viru County, Estonia*Andi, Dagestan, small village in Botlikh District, Dagestan, Russia...

, the first transgenic primate; ANDi carries foreign gene
Gene
A gene is a molecular unit of heredity of a living organism. It is a name given to some stretches of DNA and RNA that code for a type of protein or for an RNA chain that has a function in the organism. Living beings depend on genes, as they specify all proteins and functional RNA chains...

s originally from a jellyfish
Jellyfish
Jellyfish are free-swimming members of the phylum Cnidaria. Medusa is another word for jellyfish, and refers to any free-swimming jellyfish stages in the phylum Cnidaria...

.

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
University of Puerto Rico
The University of Puerto Rico is the state university system of Puerto Rico. The system consists of 11 campuses and has approximately 64,511 students and 5,300 faculty members...

 on the island of Cayo Santiago
Cayo Santiago
Cayo Santiago, also known as Isla de los monos , is an uninhabited island, located at , 950 m southeast of Punta Santiago, Humacao, Puerto Rico.-Geography:...

, off Puerto Rico
Puerto Rico
Puerto Rico , officially the Commonwealth of Puerto Rico , is an unincorporated territory of the United States, located in the northeastern Caribbean, east of the Dominican Republic and west of both the United States Virgin Islands and the British Virgin Islands.Puerto Rico comprises an...

. There are no predators 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 50% of its food comes from natural foraging. In other more controlled settings, these macaques often enjoy Fig Newton
Fig Newton
The Fig Newton is a Nabisco trademarked version of the ancient fig roll pastry filled with fig paste. Their unusual shape is a characteristic that has been adopted by many competitors including generic fig bars sold by most supermarkets.-History:...

s, and are particularly keen on "pouching" large quantities of marshmallow
Marshmallow
The marshmallow is a confection that, in its modern form, typically consists of sugar, corn syrup, water, gelatin that has been softened in hot water, dextrose, vanilla flavourings, and sometimes colouring, whipped to a spongy consistency. Some marshmallow recipes call for egg whites...

.

Sequencing the genome

Work on the genome
Genome
In modern molecular biology and genetics, the genome is the entirety of an organism's hereditary information. It is encoded either in DNA or, for many types of virus, in RNA. The genome includes both the genes and the non-coding sequences of the DNA/RNA....

 of the Rhesus macaque was completed in 2007, making Rhesus macaque the second non-human primate
Primate
A primate is a mammal of the order Primates , which contains prosimians and simians. Primates arose from ancestors that lived in the trees of tropical forests; many primate characteristics represent adaptations to life in this challenging three-dimensional environment...

 to have its genome sequenced. Humans and macaques apparently share about 93% of their DNA sequence
DNA sequence
The sequence or primary structure of a nucleic acid is the composition of atoms that make up the nucleic acid and the chemical bonds that bond those atoms. Because nucleic acids, such as DNA and RNA, are unbranched polymers, this specification is equivalent to specifying the sequence of...

 and shared a common ancestor roughly 25 million years ago.

Comparison of Rhesus macaques, chimpanzee
Chimpanzee
Chimpanzee, sometimes colloquially chimp, is the common name for the two extant species of ape in the genus Pan. The Congo River forms the boundary between the native habitat of the two species:...

s 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
Bruce Lahn is a geneticist at the University of Chicago. He is also the founder of the Center for Stem Cell Biology and Tissue Engineering at Sun Yat-sen University, Guangzhou, China. His previous research specialized in human genetics and evolutionary genetics, especially human sex chromosome...

, 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, 3 genes could be compared. If 2 genes were the same, they are 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 genome, which diverged 25 million years ago and had 93% identity, further identified evolutionary pressure and gene function.

Like the chimpanzee, changes were on the level of gene rearrangements rather than single mutations. There were frequent insertions, deletions, changes in the order and number of genes, and segmental duplications near gaps, centromeres and telomeres. So macaque, chimpanzee and human chromosomes are mosaics of each other.

Surprisingly, 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
Phenylketonuria is an autosomal recessive metabolic genetic disorder characterized by a mutation in the gene for the hepatic enzyme phenylalanine hydroxylase , rendering it nonfunctional. This enzyme is necessary to metabolize the amino acid phenylalanine to the amino acid tyrosine...

 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 3 primate species, while some are under expansion uniquely in human, chimpanzee or macaque.

For example, cholesterol pathways are conserved in all 3 species (and other primate species). In all 3 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 3 species, possibly because of climate change or mate selection. The X chromosome has 3 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
Fruit
In broad terms, a fruit is a structure of a plant that contains its seeds.The term has different meanings dependent on context. In non-technical usage, such as food preparation, fruit normally means the fleshy seed-associated structures of certain plants that are sweet and edible in the raw state,...

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 4 great ape species. The macaque genome has 33 major histocompatibility genes, 3 times that of human. This has clinical significance because the macaque is used as an experimental model of the human immune system.

In humans, the PRAME (preferentially expressed antigen of melanoma) gene family is expanded. It is actively expressed in cancers but normally testis-specific, possibly involved in spermatogenesis. The PRAME family has 26 members on human chromosome 1. In the macaque, it has 8, 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.

Agilent and Affymetrix have macaque DNA microarrays with 20,000 gene sequences, and they are used in macaque research. For example, Michael Katze of University of Washington, Seattle, infected macaques with 1918 and modern influenza. 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 7 days.

External links

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