Dueto 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, smallpox, and polio and antiretroviral medication to treat HIV/AIDS. 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 in the IUCN Red List in view of its wide distribution, presumed large population, and its tolerance of a broad range of habitats.
The name "rhesus" is reminiscent of the mythological king Rhesus of Thrace, a minor character in the Iliad. However, the French naturalist Jean-Baptiste Audebert, who applied the name to the species, stated: "it has no meaning".[4] The rhesus macaque is also known colloquially as the "rhesus monkey".[5][6]
According to Zimmermann's first description of 1780, the rhesus macaque is distributed in eastern Afghanistan, Bangladesh, Bhutan, as far east as the Brahmaputra Valley, Barak valley and in peninsular India, Nepal, and northern Pakistan. Today, this is known as the Indian rhesus macaque M. m. mulatta, which includes the morphologically similar M. rhesus villosus, described by True in 1894, from Kashmir, and M. m. mcmahoni, described by Pocock in 1932 from Kootai, Pakistan. Several Chinese subspecies 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.[7]
Rhesus macaques are native to India, Bangladesh, Pakistan, Nepal, Myanmar, Thailand, Afghanistan, Vietnam, southern China, 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 2,500 m (8,200 ft) in elevation. They are strong swimmers,[14] and can swim across rivers.[15] Rhesus macaques are noted for their tendency to move from rural to urban areas, coming to rely on handouts or refuse from humans.[16] They adapt well to human presence, and form larger troops in human-dominated landscapes than in forests.[17] 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.[18]
Kumar et al (2013)[20] 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.[20]
Fossilized isolated teeth and mandible fragments from Tianyuan Cave and a juvenile maxilla from Wanglaopu Cave near Zhoukoudian 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 decades ago is believed to have originated from Pleistocene ancestors rather than being human-introduced.[21] Fossil mandible fragments from the Taedong River Basin around Pyongyang, North Korea, have also been assigned to this species.[22]
Rhesus macaques have also been introduced to other areas, such as the United States, and become feral. The most common area for release has been Florida, with wild ranging other colonies in Puerto Rico and a semi-captive colony established in South Carolina.
Around the spring of 1938, a colony of rhesus macaques was released in and around Silver Springs in Florida by a tour boat operator known locally as "Colonel Tooey" to enhance his "Jungle Cruise". Tooey had been hoping to profit from the boom in jungle adventure stories in film and print media, buying the monkeys to be attractions at his river boat tour. Tooey apparently hadn't been aware of rhesus macaques being proficient swimmers, meaning his original plan to keep the monkeys isolated to an island inside the river didn't work. The macaques nevertheless remained in the region thanks to daily feedings by Tooey and the boat tours. Tooey subsequently released additional monkeys to add to the gene pool and avoid Inbreeding. The traditional story that the monkeys were released for scenery enhancement in the Tarzan movies that were filmed at that location is false, as the only Tarzan movie filmed in the area, 1939's Tarzan Finds a Son!, does not contain rhesus macaques.[23] Whilst this was the first colony established and the longest lasting, other colonies have since been established intentionally or accidentally. A population in Titusville, Florida was featured at the now defunct Tropical Wonderland theme park, which coincidentally was at one time endorsed by Johnny Weissmuller, who had portrayed Tarzan in the aforementioned films. This association might have contributed to the misconception the monkeys were associated directly with the Tarzan films. This colony either escaped or was intentionally released, roaming the woods of the area for a decade. In the 1980s a trapper captured several monkeys from the Titusville population and released them in the Silver Springs area to join that population.The last printed records of monkeys in the Titusville area occurred in early 1990s, but sightings continue to this day.[24]
Despite the risks, the macaques have continued to enjoy long-standing support from residents in Florida, strongly disagreeing with their removal.[24] The Silver Springs colony has continued to grow in size and range, being commonly sighted in both the park grounds, the nearby city of Ocala, Florida, and the neighboring Ocala National Forest.[28] Individuals likely originating from this colony have been seen hundreds of kilometers away, in St. Augustine, Florida and St. Petersburg, Florida. One infamous individual, titled the "Mystery Monkey of Tampa Bay", evaded capture for years in the Tampa Bay area, inspiring social media posts and a song to be written about it.[29]
Feral colonies have also resulted from research activities. There is a colony of rhesus macaques on Morgan Island, one of the Sea Islands in the South Carolina Lowcountry. They were imported in the 1970s for use in the local labs.[30][31] Another research colony was established by the Caribbean Primate Research Center of the University of Puerto Rico on the island of Cayo Santiago,[32] off of Puerto Rico. 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 in 1966. As of 2022[update] they are continuing to do ecological harm, damage crops amounting to $300,000/year and cost $1,000,000/year to manage.[33]
The Rhesus macaque is diurnal, and both arboreal and terrestrial. It is quadrupedal and, when on the ground, it walks digitigrade and plantigrade. It is mostly herbivorous, feeding mainly on fruit, but also eating seeds, roots, buds, bark, and cereals. It is estimated to consume around 99 different plant species in 46 families. During the monsoon season, it gets much of its water from ripe and succulent fruit. Rhesus macaques living far from water sources lick dewdrops from leaves and drink rainwater accumulated in tree hollows.[34] They have also been observed eating termites, grasshoppers, ants, and beetles.[35] 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.[36] Rhesus macaques have specialized pouch-like cheeks, allowing them to temporarily hoard their food.[37]It has specialised cheek pouches where it can temporarily store food and also eats invertebrates, including adult and larval insects, spiders, lice, honeycombs, crabs and bird eggs. With an increase in anthropogenic land changes, the rhesus macaque has evolved alongside intense and rapid environmental disturbance associated with human agriculture and urbanization resulting in proportions of their diet to be altered.[38]
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.[39][40] They have even been shown to demonstrate self-agency,[41] 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.[42]
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.[36] 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.[47] Rhesus social behaviour has been described as despotic, in that high-ranking individuals often show little tolerance, and frequently become aggressive towards non-kin.[48] Top-ranking female rhesus monkeys are known to sexually coerce unreceptive males and also physically injure them, biting off digits and damaging their genitals.[49]
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