View source for Rhesus macaque

{{short description|Species of Old World monkey}}
{{Speciesbox
| name = Rhesus macaque
| image = Rhesus macaque (Macaca mulatta mulatta), male, Gokarna.jpg
| image_caption = Male, Gokarna Forest, [[Nepal]]
| image2 = Rhesus Macaque in Andhra Pradesh, India.jpeg
| image2_caption = Female with infant in Andhra Pradesh
| status = LC
| status_system = IUCN3.1
| status_ref = <ref name=iucn>{{cite iucn |title=''Macaca mulatta'' |author=Singh, M. |author2=Kumar, A. |author3=Kumara, H.N. |date=2020 |article-number=e.T12554A17950825 |doi=10.2305/IUCN.UK.2020-2.RLTS.T12554A17950825.en |access-date=10 January 2022}}</ref>
| genus = Macaca
| species = mulatta
| authority = ([[Eberhard August Wilhelm von Zimmermann|Zimmermann]], 1780)<ref name=msw3>{{MSW3 Groves|pages=163|id=12100551 |heading=Species ''Macaca mulatta''}}</ref>
| range_map = Rhesus Macaque area.png
| range_map_caption = Rhesus macaque native range
| synonyms = {{hidden begin|title=Species synonymy}}
*''Simia fulvus'' <small>([[John Graham Kerr|Kerr]], 1792)</small>
*''Simia rhesus'' <small>[[Jean Baptiste Audebert|Audebert]], 1798</small>
*''Simia erythraea'' <small>[[George Shaw (biologist)|Shaw]], 1800</small>
*''Macaca nipalensis'' <small>[[Brian Houghton Hodgson|Hodgson]], 1840</small>
*''Macaca oinops'' <small>Hodgson, 1840</small>
*''Inuus sanctijohannis'' <small>[[Robert Swinhoe|R. Swinhoe]], 1866</small>
*''Inuus sancti-johannis'' <small>R. Swinhoe, 1866</small>
*''Macacus lasiotus'' <small>[[John Edward Gray|Gray]], 1868</small>
*''Macacus tcheliensis'' <small>[[Alphonse Milne-Edwards|A. Milne-Edwards]], 1872</small>
*''Macacus vestitus'' <small>A. Milne-Edwards, 1892</small>
*''Macacus rhesus villosus'' <small>[[Frederick W. True|True]], 1894</small>
*''Pithecus littoralis'' <small>[[Daniel Giraud Elliot|Elliot]], 1909</small>
*''Macaca siamica'' <small>[[Cecil Boden Kloss|Kloss]], 1917</small>
*''Macaca mulatta mcmahoni'' <small>[[Reginald Innes Pocock|Pocock]], 1932</small>
{{hidden end}}
| synonyms_ref = <ref>{{ITIS|id= 180099|taxon=''Macaca mulatta''}}</ref>
}}
[[File:Macaca mulatta 3d scan Natural History Museum University of Pisa C 1549.stl|thumb|3d model of skeleton]]
The '''rhesus macaque''' ('''''Macaca mulatta'''''), colloquially '''rhesus monkey''', is a species of [[Old World monkey]] in the ''[[Macaca]]'' genus. There are between six and nine recognised subspecies split between two groups, the Chinese-derived and the Indian-derived. Generally brown or grey in colour, it is {{cvt|47–53|cm}} in length with a {{cvt|20.7–22.9|cm}} tail and weighs {{cvt|5.3–7.7|kg}}. It is native to [[South Asia|South]], [[Central Asia|Central]], and [[Southeast Asia]] and has the widest geographic range of all non-human [[primate]]s, occupying a great diversity of altitudes and habitats. 

The rhesus macaque is [[diurnality|diurnal]], arboreal, and terrestrial. It is mostly [[herbivorous]], feeding mainly on [[fruit]], but also eating [[seed]]s, [[root]]s, [[buds]], [[Bark (botany)|bark]], and [[cereal]]s. Rhesus macaques living in cities also eat human food and trash. They are [[gregarious]], with troops comprising 20–200 individuals. The social groups are [[matrilineal]]. Individuals communicate with a variety of facial expressions, vocalisations, body postures, and gestures. 

As a result of the rhesus macaque's relatively easy upkeep, wide availability, and closeness to humans anatomically and physiologically, it has been used extensively in medical and biological research. It has facilitated many scientific breakthroughs including vaccines for [[rabies]], [[smallpox]], [[polio]], and [[antiretroviral medication]] to treat [[HIV/AIDS]]. A rhesus macaque became the first [[monkeys and apes in space|primate astronaut]] in 1948.

The rhesus is listed as [[Least Concern]] in the [[IUCN Red List]].

== Etymology ==
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 named the species, stated: "it has no meaning".<ref>{{cite book |last=Jaeger |first=E. |year=1972 |title=A source-book of biological names and terms |location=Springfield, Illinois |publisher=Charles C. Thomas}}</ref> The rhesus macaque is also known colloquially as the "rhesus monkey".<ref>{{cite web |title=Rhesus macaque monkey (1932) |publisher=[[British Society for Immunology]] |url=https://www.immunology.org/rhesus-macaque-monkey-1932 |archive-url=https://web.archive.org/web/20201108003247/https://www.immunology.org/rhesus-macaque-monkey-1932 |archive-date=2020-11-08
}}</ref><ref>{{cite news |title=Frequently asked questions about nonhuman primates in research |newspaper=National Primate Research Centers |date=
|url=https://nprcresearch.org/primate/faq.php|archive-url=https://web.archive.org/web/20151123053404/https://nprcresearch.org/primate/faq.php |archive-date=2015-11-23}}</ref>

== Taxonomy ==
[[File:Macaque India 3.jpg|thumb|Rhesus macaque by the [[Agra Fort]], [[Uttar Pradesh]], India]]
[[File:Rhesus Macaque2.jpg|thumb|Mother and child rhesus macaque in Nepal]]

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 ''Macaca mulatta mulatta'', which includes the morphologically similar ''M. rhesus villosus'', described by [[Frederick W. True|True]] in 1894, from [[Kashmir]], and ''M. m. mcmahoni'', described by [[Reginald Innes Pocock|Pocock]] in 1932 from Kootai, [[Pakistan]]. Several [[China|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.<ref name="apc04">{{cite book |author=Brandon-Jones, D. |author2=Eudey, A. A. |author3=Geissmann, T. |author4=Groves, C. P. |author5=Melnick, D. J. |author6=Morales, J. C. |author7=Shekelle, M. |author8=Stewart, C.-B. |title=Asian primate classification |publisher= International Journal of Primatology |volume=25 |issue=1 |date=2004 |pages=97–164 |url=http://www.gibbons.de/main/papers/pdf_files/2004asianprimates.pdf |access-date=5 January 2016}}</ref>

The Chinese subspecies can be divided as follows:
* ''M. m. mulatta'' is found in western and central China, in the south of [[Yunnan]], and southwest of [[Guangxi]];<ref name="shilai1991">{{cite journal |author1=Jiang, X. |author2=Wang, Y. |author3=Ma, S. |year=1991 |title=Taxonomic revision and distribution of subspecies of rhesus monkey (''Macaca mulatta'') in China |journal=Zoological Research |volume=12 |url=http://www.zoores.ac.cn/EN/abstract/abstract1901.shtml}}</ref>
* ''M. m. lasiota'' ([[John Edward Gray|Gray]], 1868), the west Chinese rhesus macaque, is distributed in the west of [[Sichuan]], northwest of [[Yunnan]], and southeast of [[Qinghai]];<ref name="shilai1991" /> it is possibly synonymous with ''M. m. sanctijohannis'' ([[Robert Swinhoe|R. Swinhoe]], 1867), if not with ''M. m. mulatta''.<ref name="apc04" />
* ''M. m. tcheliensis'' ([[Henri Milne-Edwards|Milne-Edwards]], 1870), the north Chinese rhesus macaque, lives in the north of [[Henan]], south of [[Shanxi]], and near [[Beijing]]. Some consider it as the most endangered subspecies.<ref>{{cite journal |doi=10.1007/BF02215449 |title=Phylogeny of rheusus monkeys (''Macaca mulatta'') as revealed by mitochondrial DNA restriction enzyme analysis |journal=International Journal of Primatology |volume=14 |issue=4 |page=587 |year=1993 |last1=Zhang |first1=Y. |last2=Shi |first2=L. |s2cid=21883524}}
</ref> Others consider it possibly synonymous with ''M. m. sanctijohannis'', if not with ''M. m. mulatta''.<ref name="apc04" />
* ''M. m. vestita'' ([[Henri Milne-Edwards|Milne-Edwards]], 1892), the Tibetan rhesus macaque, lives in the southeast of [[Tibet]], northwest of Yunnan ([[Deqin County|Deqing]]), and perhaps including [[Yushu Tibetan Autonomous Prefecture|Yushu]];<ref name="shilai1991" /> it is possibly synonymous with ''M. m. sanctijohannis'', if not with ''M. m. mulatta''.<ref name="apc04" />
* ''M. m. littoralis'' ([[Daniel Giraud Elliot|Elliot]], 1909), the south Chinese rhesus macaque, lives in [[Fujian]], [[Zhejiang]], [[Anhui]], [[Jiangxi]], [[Hunan]], [[Hubei]], [[Guizhou]], northwest of [[Guangdong]], north of [[Guangxi]], northeast of [[Yunnan]], east of [[Sichuan]], and south of [[Shaanxi]];<ref name="shilai1991" /> it is possibly synonymous with ''M. m. sanctijohannis'', if not with ''M. m. mulatta''.<ref name="apc04" />
* ''M. m. brevicaudus'', also referred to as ''Pithecus brevicaudus'' ([[Daniel Giraud Elliot|Elliot]], 1913), lives on the [[Hainan Island]] and [[Wanshan Islands]] in [[Guangdong]], and the islands near [[Hong Kong]];<ref name="shilai1991" /> it may be synonymous with ''M. m. mulatta''.<ref name="apc04" />
* ''M. m. siamica'' ([[Cecil Boden Kloss|Kloss]], 1917), the Indochinese rhesus macaque, is distributed in [[Myanmar]], in the north of [[Thailand]] and [[Vietnam]], in [[Laos]], and in the Chinese provinces of [[Anhui]], northwest [[Guangxi]], Guizhou, [[Hubei]], [[Hunan]], central and eastern [[Sichuan]], and western and south-central [[Yunnan]]; possibly synonymous with ''M. m. sanctijohannis'', if not with ''M. m. mulatta''.<ref name="apc04" />

== Description ==
[[File:Rhesus Macaque, Red Fort, Agra.jpg|thumb|Male Rhesus macaque in Agra fort, Uttar Pradesh]]
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 {{convert|20.7|and|22.9|cm|in|abbr=on}}.<ref name="Fooden2000">{{cite journal |doi=10.5962/bhl.title.7192 |title=Systematic review of the rhesus macaque, ''Macaca mulatta'' (Zimmermann, 1780) |year=2000 |last1=Fooden |first1=J. |journal=Fieldiana |volume=96 |pages=1–180|doi-access=free }}</ref> Adult males measure about {{convert|53|cm|in|abbr=on}} on average and weigh about {{convert|7.7|kg|lb|abbr=on}}. Females are smaller, averaging {{convert|47|cm|in|abbr=on}} in length and {{convert|5.3|kg|lb|abbr=on}} in weight.<ref name="Fooden2000" /><ref name="SinghSinha2004">{{cite book |last1=Singh |first1=M. |last2=Sinha |first2=A. |year=2004 |chapter=Life history traits: ecological adaptations or phylogenetic relics? |editor1-last=Thierry |editor1-first=B. |editor2-last=Singh |editor2-first=M. |editor3-last=Kaumanns |editor3-first=W. |title=Macaque societies: a model for the study of social organization |location=Cambridge, UK |publisher=Cambridge University Press |pages=80–83 |isbn=978-0-521-52168-0}}</ref> The [[intermembral index|ratio of arm length to leg length]] is 89.6–94.3%.<ref name="Hamada_etal2005">{{cite journal |last1=Hamada |first1=Y. |last2=Watanabe |first2=T. |last3=Chatani |first3=K. |last4=Hayakawa |first4=S. |last5=Iwamoto |first5=M. |year=2005 | title=Morphometrical comparison between Indian- and Chinese-derived rhesus macaques (''Macaca mulatta'') |journal=Anthropological Science |volume=113 |issue=2 |pages=183–188 |doi=10.1537/ase.03104 |doi-access=free}}</ref>

The rhesus macaque has a dental formula of {{DentalFormula|upper=2.1.2.3|lower=2.1.2.3|total=32}} and [[bilophodont]] [[Molar (tooth)|molar teeth]].<ref>{{cite book |author=Pocock, R. I. |year=1939 |title=The fauna of British India, including Ceylon and Burma: Mammalia volume I |location=London |publisher=Taylor and Francis |pages=15–185 |chapter=Catarhini |chapter-url=https://archive.org/stream/PocockMammalia1/pocock1#page/n49/mode/2up}}</ref>

== Distribution and habitat ==
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 {{convert|2500|m|ft|abbr=on}} in elevation. They are strong swimmers,<ref>{{cite book |doi=10.1016/B978-0-12-417144-2.00006-8 |chapter=Basic physiology of ''Macaca mulatta'' |title=The nonhuman primate in nonclinical drug development and safety assessment |year=2015 |last1=Lewis |first1=A. D. |last2=Prongay |first2=K. |pages=87–113 |isbn=978-0-12-417144-2}}</ref> and can swim across rivers.<ref name="Anderson_etal2016">{{cite report |publisher=Department of Wildlife Ecology and Conservation, UF/IFAS Extension |year=2016 |title=History and status of introduced rhesus macaques (''Macaca mulatta'') in Silver Springs State Park, Florida |first1=C. J. |last1=Anderson |first2=S. A. |last2=Johnson |first3=M. E. |last3=Hostetler |first4=M. G. |last4=Summers |url=https://edis.ifas.ufl.edu/publication/uw412}}</ref> Rhesus macaques are noted for their tendency to move from rural to urban areas, coming to rely on handouts or refuse from humans.<ref>{{cite journal |author=Ciani, A. C. |year=1986 |title=Intertroop agonistic behavior of a feral rhesus macaque troop in ranging in town and forest areas in India |journal=Aggressive Behavior |volume=12 |issue=6 |doi=10.1002/1098-2337(1986)12:6<433::AID-AB2480120606>3.0.CO;2-C |pages=433–439 |url=http://www.esploratore.com/files/resourcesmodule/@random44be4ac49f856/1153323855_Intertroop_Agonistic_Behavior_of_a_Feral_rhesus_Macaque_Troop_Ranging_in_Town_and_Areas_in_India.pdf |access-date=5 January 2016}}</ref> They adapt well to human presence, and form larger troops in human-dominated landscapes than in forests.<ref>{{cite journal |first1=K. |last1=Rishi |first2=S. |last2=Anindya |first3=R. |last3=Sindhu |year=2013 |title=Comparative demography of two commensal macaques in India: implications for population status and conservation |journal=Folia Primatologica |volume=84 |issue=6 |pages=384–393 |doi=10.1159/000351935 |pmid=24022675 |s2cid=45152033}}</ref> 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.<ref name=":4">{{Cite journal |last1=Ji |first1=Y. |last2=Wei |first2=X. |last3=Liu |first3=F. |last4=Li |first4=D. |last5=Li |first5=J. |date=2022 |title=Spatial-Temporal Patterns of Human-Wildlife Conflicts Under Coupled Impact of Natural and Anthropogenic Factors in Mt. Gaoligong, Western Yunnan, China |journal=Global Ecology and Conservation|volume=40|article-number=e02329|doi=10.1016/j.gecco.2022.e02329 |bibcode=2022GEcoC..4002329J |doi-access=free}}</ref>

The southern and the northern distributional limits for rhesus and [[bonnet macaque]]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.<ref>{{cite journal |last1=Kumar |first1=R. R. |last2=Sinha |first2=A. |year=2011 |title=Of least concern? Range extension by rhesus macaques (''Macaca mulatta'') threatens long-term survival of bonnet macaques (''M. radiata'') in Peninsular India |url=https://nias.academia.edu/RishiKumar/Papers/538383/Of_least_concern_Range_Extension_by_Rhesus_Macaques_Macaca_mulatta_Threatens_Long-Term_Survival_of_Bonnet_Macaques_M._radiata_in_Peninsular_India |journal=International Journal of Primatology |volume=32 |issue=4 |pages=945–959 |doi=10.1007/s10764-011-9514-y |s2cid=35227050 |access-date=5 January 2016}}</ref>

Kumar et al (2013)<ref name=":04">{{Cite journal |last1=Kumar |first1=R. |last2=Sinha |first2=A. |last3=Radhakrishna |first3=S. |date=2013 |title=Comparative Demography of Two Commensal Macaques in India: Implications for Population Status and Conservation |url=https://www.researchgate.net/publication/256490353 |journal=Folia Primatologica |volume=84 |issue=6 |pages=384–393 |doi=10.1159/000351935 |pmid=24022675 |s2cid=45152033}}</ref> 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.<ref name=":04" />

===Fossil record===
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.<ref>{{Cite journal |last=Tong |first=H. |date=2014 |title=Fossils of Macaca mulatta from Tianyuan Cave, with Notes on Some Previously Unpublished Specimens of Macaca from Zhoukoudian Area |journal=Acta Geologica Sinica |volume=88 |issue=5 |pages=1397–1408 |doi=10.1111/1755-6724.12307|bibcode=2014AcGlS..88.1397T |s2cid=128707124}}</ref> Fossil [[mandible]] fragments from the [[Taedong River]] Basin around [[Pyongyang]], North Korea, have also been assigned to this species.<ref>{{Cite journal |last1=Han |first1=K. S. |last2=So |first2=K. S. |last3=Choe |first3=R. S. |last4=Kang |first4=J. C. |last5=Kang |first5=I. |last6=Ri |first6=C. U. |date=2022 |title=First record of ''Macaca mulatta'' (Cercopithecidae: Papionini) from the Taedong River Basin, the Democratic People's Republic of Korea |journal=Palaeoworld |volume=32 |issue=3 |pages=573–578 |doi=10.1016/j.palwor.2022.09.008 |s2cid=252768097}}</ref>

=== [[Exogeny|Exogenous]] colonies ===
Rhesus macaques have also been introduced and acclimated to other areas, such as the United States, where they are considered an invasive species.<ref>{{Cite news |date=1997 |title=Monkey Business: For 25 years, furry residents of tiny islands have been real swingers |url=https://www.newspapers.com/image/990297062/?match=1&terms=%22outwitted%20by%20monkeys%22 |access-date=November 12, 2024 |work=AP}}</ref>
Around the spring of 1938, a colony of rhesus macaques was released in and around [[Silver Springs (attraction)|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.<ref>{{cite book |title=Primates face to face |last=Wolfe |first=L. |publisher=Cambridge University Press |year=2002 |isbn=0-521-79109-X |page=320 |url=https://books.google.com/books?id=zYg3eOqjq4UC&pg=PA320}}</ref> 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.<ref name="Anderson et al">{{cite journal |last1=Anderson |first1=C. J. |last2=Hostetler |first2=M. E. |last3=Johnson |first3=A. |title=History and Status of Introduced Non-Human Primate Populations in Florida |journal=Southeastern Naturalist |date=2017 |volume=16 |issue=1 |pages=19–36 |doi=10.1656/058.016.0103}}</ref>

Various colonies of rhesus macaque are speculated to be the result of [[zoos]] and wildlife parks destroyed in hurricanes, most notably [[Hurricane Andrew]].<ref>{{cite news |url=http://www.newyorker.com/reporting/2009/04/20/090420fa_fact_bilger |author=Bilger, B. |title=The natural world, swamp things |newspaper=The New Yorker |date= 20 April 2009 |page=80 |access-date=5 January 2016}}</ref> A 2020 estimate put the number at 550–600 rhesus macaques living in the state;<ref>{{cite news |url=https://www.washingtonpost.com/science/2020/02/25/rhesus-macaques-herpes-florida/ |title=People can't agree on what to do about Florida's herpes-infected monkeys |first=Hannah |last=Knowles
|date=25 February 2020 |access-date=5 September 2021 |newspaper=[[The Washington Post]]}}</ref> officials have caught more than 1,000 of the monkeys in the past decade. Most of the captured monkeys tested positive for [[herpes B virus]], which leads wildlife officials to consider the animals a public health hazard.<ref>{{cite news |url=https://nypost.com/2013/09/12/herpes-infected-monkeys-terrorize-florida/ |title=Herpes-infected monkeys terrorize Florida |work=[[New York Post]] |first=Natalie |last=O'Neill |date=12 September 2013}}</ref>

Despite the risks, the macaques have continued to enjoy long-standing support from residents in Florida, strongly disagreeing with their removal.<ref name="Anderson et al" /> 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]].<ref>{{cite web |title=The Silver Springs monkeys - International Primate Protection League |url=https://ippl.org/blog/the-silver-springs-monkeys/ |website=International Primate Protection League |access-date=13 May 2023 |date=2013}}</ref> Individuals likely originating from this colony have been seen hundreds of kilometers away, in [[St. Augustine, Florida]] and [[St. Petersburg, Florida]]. One infamous individual, named the "Mystery Monkey of [[Tampa Bay area|Tampa Bay]]", evaded capture for years, inspiring social media posts and a song.<ref>{{cite web |last1=Hayes |first1=S. |title=Where is the Mystery Monkey of Tampa Bay? |url=https://www.tampabay.com/special-reports/2021/11/09/where-is-the-mystery-monkey-of-tampa-bay/ |work=Tampa Bay Times}}</ref>

Exogenous colonies have also resulted from research activities. There is a colony of rhesus macaques on [[Morgan Island, South Carolina|Morgan Island]], one of the [[Sea Islands]] in the [[South Carolina Lowcountry]]. They were imported in the 1970s for use in the local labs.<ref>[http://www.thestate.com/mld/thestate/news/local/8961941.htm The State | Homepage<!-- Bot generated title -->] {{webarchive|url=https://web.archive.org/web/20040826195106/http://www.thestate.com/mld/thestate/news/local/8961941.htm |date=2004}}</ref><ref>{{cite journal |author1=Taub, D.M. |author2=Mehlman, P.T. |title=Development of the Morgan Island rhesus monkey colony |journal=P R Health Sci J |volume=8 |issue=1 |pages=159–69 |date=1989 |pmid=2780958}}</ref> Another research colony was established by the Caribbean Primate Research Center of the [[University of Puerto Rico]] on the island of [[Cayo Santiago]],<ref name="Rawlins-1986">{{cite book |last1=Rawlins | first1=R. G. | last2=Kessler |first2=M. J. |title=The Cayo Santiago Macaques: History, Behavior, and Biology |series=SUNY series in primatology |publisher=State University of New York Press |place=Albany |year=1986 |isbn=978-0-88706-135-6 | oclc=42855829 |page=306}}</ref> 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}} they are continuing to do ecological harm, damage crops amounting to $300,000/year and cost $1,000,000/year to manage.<ref name="UFLIFASEDIS">{{cite web | title=WEC367/UW412: History and Status of Introduced Rhesus Macaques (''Macaca mulatta'') in Silver Springs State Park, Florida | website=Ask IFAS, Electronic Data Information Source (EDIS) | publisher=[[Institute of Food and Agricultural Sciences]] (IFAS), UF | date=2022 | url=http://edis.ifas.ufl.edu/publication/UW412 | access-date=2022-02-09}}</ref>

== Ecology and behavior ==
[[File:Macaque Haji Malang.jpg|thumb|Rhesus macaque displaying its canine teeth]]
The Rhesus macaque is [[Diurnality|diurnal]], and both [[arboreal]] and [[Terrestrial animal|terrestrial]]. It is [[quadruped]]al and, when on the ground, it walks [[digitigrade]] and [[plantigrade]]. It is mostly [[herbivorous]], feeding mainly on [[fruit]], but also eating [[seed]]s, [[root]]s, [[buds]], [[Bark (botany)|bark]], and [[cereal]]s. 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.<ref name="Makwana 1979">{{cite journal |last1=Makwana |first1=S. |year=1979 |title=Field ecology and behavior of the rhesus macaque. Food, feeding and drinking in Dehra Dun forests |journal=Indian Journal of Forestry |volume=2 |issue=3 |pages=242–253}}</ref> They have also been observed eating [[termite]]s, [[grasshopper]]s, [[ant]]s, and [[beetle]]s.<ref>{{cite book |last=Lindburg |first=D. G. |year=1971 |chapter=The rhesus monkeys in north India: an ecological and behavioural study |editor-last=Rosenblum |editor-first=L. A. |title=Primate behaviour: developments in the field and laboratory research |publisher=Academic Press |location=New York |volume=1 |pages=83–104 |isbn=0-12-534001-X}}</ref> 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.<ref name="Southwick 1965"/> Rhesus macaques have specialized pouch-like cheeks, allowing them to temporarily hoard their food.<ref name="Burrows-et-al-2009">{{cite journal |last1=Burrows |first1=A. M. |last2=Waller | first2=B. M. |last3=Parr |first3=L. A. |title=Facial musculature in the rhesus macaque (''Macaca mulatta''): evolutionary and functional contexts with comparisons to chimpanzees and humans |journal=Journal of Anatomy |volume=215 |issue=3 |year=2009 |issn=0021-8782 |doi=10.1111/j.1469-7580.2009.01113.x |pages=320–334 |pmid=19563473 |pmc=2750044}}</ref>
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.<ref name=Cooper2022>{{Cite journal |last1=Cooper |first1=E. B. |last2=Brent |first2=L. J. N. |last3=Snyder-Mackler |first3=N. |last4=Singh |first4=M. |last5=Sengupta |first5=A. |last6=Khatiwada |first6=S. |last7=Malaivijitnond |first7=S. |last8=Qi Hai |first8=Z. |last9=Higham |first9=J. P. |date=2022 |title=The rhesus macaque as a success story of the Anthropocene |journal=eLife |volume=11 |article-number=e78169 |doi=10.7554/eLife.78169 |pmc=9345599 |pmid=35801697 |doi-access=free}}</ref>

In [[comparative psychology|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.<ref>{{Cite journal |author=Couchman, J. J. |title=Beyond stimulus cues and reinforcement signals: a new approach to animal metacognition |journal=Journal of Comparative Psychology |volume=124 |issue=4 |pages=356–368 |year=2010 |doi=10.1037/a0020129 |pmid=20836592 |pmc=2991470 |display-authors=etal}}</ref><ref>{{cite journal |last1=Blanchard |first1=T. C. |last2=Wolfe |first2=L. S. |last3=Vlaev |first3=I. |last4=Winston |first4=J. S. |last5=Hayden |first5=B. Y. |title=Biases in preferences for sequences of outcomes in monkeys |journal=Cognition |volume=130 |issue=3 |pages=289–299 |doi=10.1016/j.cognition.2013.11.012 |year=2014 |pmid=24374208 |pmc=3969290}}</ref> They have even been shown to demonstrate [[self-agency]],<ref>{{Cite journal |author=Couchman, J. J. |title=Self-agency in rhesus monkeys |journal=Biology Letters |year=2011 |doi=10.1098/rsbl.2011.0536 |volume=8 |pages=39–41 |pmid=21733868 |issue=1 |pmc=3259954}}</ref> 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.<ref>{{cite magazine |last=Waxman |first=O. B. |date=2014 |url=https://time.com/3643900/monkey-electrocution-india/ |title=Hero monkey revives simian pal electrocuted in India |magazine=Time}}</ref>

=== Group structure ===
[[File:Rhesus macaque monkey family D72 16866k.jpg|thumb|Rhesus macaque adult females with baby, IIT Mandi, Himachal, India. Aug '20]]
Like other macaques, rhesus troops comprise a mixture of 20–200 males and females.<ref>{{cite book |author1=Teas, J. |author2=Richie, T. |author3=Taylor, H. |author4=Southwick, C. |year=1980 |chapter=Population patterns and behavioral ecology of rhesus monkeys (''Macaca mulatta'') in Nepal |title=The macaques: studies in ecology, behavior, and evolution |editor=Lindenburg, D. |place=San Francisco |publisher=Van Nostrand Reinhold Company |isbn=0-442-24817-2}}</ref> Females may outnumber the males by a ratio of 4:1. Males and females both have separate hierarchies. Female [[philopatry]], common among social mammals, has been extensively studied in rhesus macaques. Females tend not to leave the social group, and 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 existing in a hierarchy, and a female outranks any unrelated females that rank lower than her mother.<ref>{{cite journal |author1=Judge, P.  |author2=Waal, F. |name-list-style=amp |year=1997 |title=Rhesus monkey behaviour under diverse population densities: coping with long-term crowding |journal=Animal Behaviour |volume=54 |issue=3 |pages=643–662 |doi=10.1006/anbe.1997.0469 |pmid=9299049 |s2cid=22512222}}</ref> Rhesus macaques are unusual in that the youngest females tend to outrank their older sisters.<ref>{{cite book |last=Waal |first=F. |year=1993 |chapter=Codevelopment of dominance relations and affiliative bonds in rhesus monkeys |title=Juvenile primates: life history, development, and behavior |editor1-last=Pereira |editor1-first=M. |editor2-first=L. |editor2-last=Fairbanks |location=Oxford, UK |publisher=Oxford University Press |isbn=0-226-65622-5}}</ref> This is likely because young females are more fit and fertile. Mothers seem to prevent the older daughters from forming coalitions against her.{{clarify |date = January 2016 |reason = "her" who? the mother, or the youngest daughter?}} 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.<ref name="Hill 1993">{{cite book |last1=Hill |first1=D. |last2=Okayasu |first2=N. |year=1996 |chapter=Determinants of dominance among female macaques: nepotism, demography and danger |title=Evolution and ecology of macaque societies |editor1-last=Fa |editor1-first=J. |editor2-first=D. |editor2-last=Lindburg |location=Cambridge |publisher=Cambridge University Press |isbn=0-521-41680-9}}</ref> 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.<ref name="Southwick 1965">{{cite book |last1=Southwick |first1=C. |last2=Beg |first2=M. |first3=R. |last3=Siddiqi |year=1965 |chapter=Rhesus monkeys in North India |title=Primate behavior: field studies of monkeys and apes |editor-last=DeVore |editor-first=I. |location=San Francisco, USA |publisher=Holt, Rinehart and Winston |isbn=978-0-03-050340-5}}</ref>

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.<ref name="Southwick 1965"/> 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.<ref name="Gouzoules 1998">{{cite journal |author1=Gouzoules, H. |author2=Gouzoules, S. |author3=Tomaszycki, M. |year=1998 |title=Agonistic screams and the classification of dominance relationships: are rhesus monkeys fuzzy logicians? |journal=Animal Behaviour |volume=55 |issue=1 |pages=51–60 |doi=10.1006/anbe.1997.0583 |pmid=9480671 |s2cid=25747013 }}</ref> Rhesus social behaviour has been described as despotic, in that high-ranking individuals often show little tolerance, and frequently become aggressive towards non-kin.<ref>{{cite journal |last1=Thierry |first1=B. |year=1985 |title=Social development in three species of macaque (''Macaca mulatta'', ''M. fasicularis'', ''M. tonkeana''): A preliminary report on the first ten weeks of life |journal=Behavioural Processes |volume=11 |issue= 1|pages=89–95 |doi=10.1016/0376-6357(85)90105-6 |pmid=24924364 |s2cid=140208622}}</ref> Top-ranking female rhesus monkeys are known to sexually coerce unreceptive males and also physically injure them, biting off digits and damaging their genitals.<ref>{{Cite web |title=Queen Kong? In rhesus realm, females rule |url=https://www.chicagotribune.com/news/ct-xpm-2007-11-19-0711190082-story.html |access-date=2021-02-12 |website=chicagotribune.com|date=19 November 2007 }}</ref>

Rhesus macaques have been observed engaging in interspecies grooming with [[Hanuman langur]]s and with [[Sambar deer]].<ref>{{cite journal |title=First record of interspecies grooming between Raffles' banded langur and long-tailed macaque |last1=Lee |first1=Z.H. |last2=Ang |first2=A. |last3=Ruppert |first3=N. |journal=Journal of Threatened Taxa |date=26 August 2021 |doi=10.11609/jott.7510.13.9.19246-19253 |pages=19246–19253 |volume=13 |issue=9 |url=https://threatenedtaxa.org/index.php/JoTT/article/view/7510/7884|access-date=2021-09-05|doi-access=free }}</ref>

=== 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.<ref name="Maestripieri 1999" >{{cite book |last=Maestripieri |first=D. |year=1999 |chapter=Primate social organization, gestural repertoire size, and communication dynamics: a comparative study of macaques |pages=55–77 |editor-last=King |editor-first=B. J. |title=The origins of language: what nonhuman primates can tell us |location=Santa Fe, NM, USA |publisher=School American Research Press |isbn=0-85255-904-6}}</ref> 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.<ref>{{cite book |last=Rowe |first=N. |year=1996 |title=The pictorial guide to the living primates |location=East Hampton, NY, USA |publisher=Pogonias Press}}</ref> Another submissive behavior is the "present rump", where an individual raises its tail and exposes its genitals to the dominant one.<ref name="Maestripieri 1999" /> A dominant individual threatens another individual by standing quadrupedally and making a silent "open mouth stare" accompanied by the tail sticking straight.<ref>{{cite journal |author=Partan, S. R. |year=2002 |title=Single and multichannel signal composition: facial expressions and vocalizations of rhesus macaques (''Macaca mulatta'')  |journal=Behaviour |volume=139 |issue=2–3 |pages=993–1027 |doi=10.1163/15685390260337877}}</ref> During movements, macaques make coos and grunts. These are also made during affiliative interactions, and approaches before grooming.<ref>{{cite journal |author=Hauser, M. D. |year=1998 |title=Functional referents and acoustic similarity field playback experiments with rhesus monkeys |journal=Animal Behaviour |volume=55 |issue=6 |pages=1647–58 |doi=10.1006/anbe.1997.0712 |pmid=9642008 |s2cid=8253916}}</ref> 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.<ref name="Lindburg 1971">{{cite book |last=Lindburg |first=D. G. |year=1971 |chapter=The rhesus monkey in north India: an ecological and behavioral study |pages=1–106 |editor-last=Rosenblum |editor-first=L. A. |title=Primate behavior: developments in field and laboratory research |volume=2 |location=New York |publisher=Academic Press |isbn=1-4832-4440-7}}</ref> Screeches, screams, squeaks, pant-threats, growls, and barks are used during aggressive interactions.<ref name="Lindburg 1971"/> Infants "[[gecker]]" to attract their mother's attention.<ref name="Patel">{{cite journal |doi=10.1002/ajp.20027 |title=Acoustic and behavioral analyses of 'gecker' distress vocalizations in young rhesus macaques (''Macaca mulata'') |journal=American Journal of Primatology |volume=62 |issue=5 |page=48 |year=2004 |author1=Patel, E. R. |author2=Owren, M. J. |bibcode=2004ASAJ..115.2485P}}</ref>

=== Reproduction ===
[[File:Davidraju img15(Macaca mulatta) Rhesus macaque.jpg|thumb|Mother rhesus macaque with her baby]]
Adult male macaques try to maximize their reproductive success by entering into [[sex]] 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.<ref name="Bercovitch 1997">{{cite journal |author=Bercovitch, F. |year=1997 |title=Reproductive strategies of rhesus macaques |journal=Primates |volume=38 |issue=3 |pages=247–263 |doi=10.1007/BF02381613 |s2cid=8372920}}</ref> Female macaques first breed when they are four years old and reach [[menopause]] at around twenty-five years of age.<ref name="MLWalker2">{{cite journal |author1=Walker, M. L. |author2=Herndon, J. G. |title=Menopause in nonhuman primates? |journal=Biology of Reproduction |volume=79 |pages=398–406 |year=2008 |pmid=18495681 |doi=10.1095/biolreprod.108.068536 |issue=3 |pmc=2553520}}</ref> Male macaques generally play no role in raising the young but do have peaceful relationships with the offspring of their consort pairs.<ref name="Southwick 1965"/>

Manson and Parry<ref name="pmid8460656">{{cite journal |last1=Manson |first1=J. H. |last2=Perry |first2=S. E. |title=Inbreeding avoidance in rhesus macaques: whose choice? |journal=American Journal of Physical Anthropology |volume=90 |issue=3 |pages=335–44 |year=1993 |pmid=8460656 |doi=10.1002/ajpa.1330900307 |bibcode=1993AJPA...90..335M |url=https://deepblue.lib.umich.edu/bitstream/2027.42/37662/1/1330900307_ftp.pdf |hdl=2027.42/37662 |hdl-access=free}}</ref> 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]] the preceding birth season.<ref name = "Berman 1992">{{cite journal |author=Berman, C. |year=1992 |title=Immature siblings and mother-infant relationships among free-ranging rhesus monkeys on Cayo Santiago |journal=Animal Behaviour |volume=44 |pages=247–258 |doi=10.1016/0003-3472(92)90031-4 |s2cid=53171870}}</ref> Infants farther from the center of the groups are more vulnerable to [[infanticide]] from outside groups.<ref name="Southwick 1965"/> Some mothers abuse their infants, which is believed to be the result of controlling [[parenting styles]].<ref>{{cite journal |author=Maestripieri, D. |year=1998 |title=Parenting styles of abusive mothers in group-living rhesus macaques |journal=Animal Behaviour |volume=55 |issue=1 |pages=1–11 |doi=10.1006/anbe.1997.0578 |pmid=9480666 |s2cid=8348608}}</ref>

Primatologist [[Robert Goy]] discovered that adult Rhesus macaques will take care of an unrelated infant in captivity. A male Rhesus macaque will ignore an infant as long as a female Rhesus macaque is around. If a male Rhesus macaque is the only option for parental care, they will tend to an unrelated infant like a female would.<ref>{{cite book |last1=de Waal |first1=Frans |title=Different: Gender Through the Eyes of a Primatologist |date=2022 |publisher=W.W. Norton |isbn=978-1-324-00710-4 |page=254-255}}</ref>

===Aging===
The rhesus monkey has been used as a model for studying aging of the [[ovary|ovaries]] of primate females.<ref name = Zhang2015>{{cite journal |last1=Zhang |first1=D |last2=Zhang |first2=X |last3=Zeng |first3=M |last4=Yuan |first4=J |last5=Liu |first5=M |last6=Yin |first6=Y |last7=Wu |first7=X |last8=Keefe |first8=DL |last9=Liu |first9=L. |display-authors=4 |title=Increased DNA damage and repair deficiency in granulosa cells are associated with ovarian aging in rhesus monkey |journal=Journal of Assisted Reproduction and Genetics |year=2015 |volume=32 |issue=7 |pages=1069–1078 |doi=10.1007/s10815-015-0483-5 |pmid=25957622 |pmc=4531862}}</ref> Ovarian aging was found to be associated with increased [[DNA damage (naturally occurring)|DNA double strand breaks]] and reduced [[DNA repair]] in [[granulosa cell]]s, that is, [[somatic cell]]s closely associated with developing [[oocyte]]s.<ref name = Zhang2015/>

=== Self-awareness ===
In several [[mirror test|experiments]] 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 [[Self-awareness|aware of themselves]].<ref>{{cite journal |author1=Rajala, A. Z. |author2=Reininger, K. R. |author3=Lancaster, K. M. |author4=Populin, L. C. |name-list-style=amp |year=2010 |title=Rhesus monkeys (''Macaca mulatta'') do recognize themselves in the mirror: implications for the evolution of self-recognition |journal=PLOS ONE 
|doi=10.1371/journal.pone.0012865 |volume=5 |issue=9 |article-number=e12865 |pmid=20927365 |pmc=2947497 |bibcode=2010PLoSO...512865R |doi-access=free}}</ref>

===Predators===
[[Common thresher shark]]s and [[Ganges shark]]s prey on rhesus macaques when it swim across [[estuaries]] at [[ebb tide]].<ref>{{Cite book|title=Primates as prey: ecological, morphological and behavioral relationships between primate species and their predators|date=May 2000|publisher=Washington University Department of Anthropology|author=Hart, D. L.}}</ref>

=== Human–rhesus conflict ===
The macaque–human relationships is complex and culturally specific, ranging from relatively peaceful coexistence to extreme levels of conflict.<ref name=Priston2012>{{Cite book |last1=Priston |first1=N. E. C. |last2=McLennan |first2=M. R.|doi=10.1007/978-1-4614-3967-7_14 |chapter=Managing Humans, Managing Macaques: Human–Macaque Conflict in Asia and Africa |pages=225–250 |date=2012 |title=The Macaque Connection |editor1=Radhakrishna, S. |editor2=Huffman, M. A. |editor3=Sinha, A. |place=New York, NY |publisher=Springer New York |isbn=978-1-4614-3966-0}}</ref> Conflicts tend to result from rapidly changing agricultural practices, increasing urbanisation, and clearing of woodlands and other territory, pushing macaques into human settlements in the search for resources.<ref name=":5">{{Cite journal |last1=Gopalan |first1=R. |last2=Radhakrishna |first2=S. |date=2022 |title=Moving From Coexistence to Conflict: A Political Ecology Perspective On Human-Rhesus Macaque Conflict in Himachal Pradesh, India |journal=Human Ecology |volume=50 |issue=3 |pages=463–476 |doi=10.1007/s10745-022-00331-7 |bibcode=2022HumEc..50..463G |s2cid=250156685}}</ref> A 2021 study stated that human-macaque conflict is one of the most critical challenges faced by wildlife managers in the South- and Southeast-Asian regions.<ref name=":12" /> 

Conflict between rhesus macaques and humans is at all-time high, with areas once forested habitat being converted to industrial agriculture. In Nepal, 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.<ref name=":12">{{Cite journal |last1=Koirala |first1=S. |last2=Garber |first2=P. A. |last3=Somasundaram |first3=D. |last4=Katuwal |first4=H. B. |last5=Ren |first5=B. |last6=Huang |first6=C. |last7=Li |first7=M. |date=2021 |title=Factors affecting the crop raiding behavior of wild rhesus macaques in Nepal: Implications for wildlife management. |journal=Journal of Environmental Management |volume=297 |article-number=113331 |doi=10.1016/j.jenvman.2021.113331 |pmid=34298347 |bibcode=2021JEnvM.29713331K}}</ref> Crop raiding is one of the biggest visible effects of human-rhesus conflict. 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.<ref name=":12" /> This has resulted in farmers and other members of the population viewing macaques inhabiting agricultural landscapes as serious crop pests.<ref name=":12" /> Nepal is a significant study area with almost 44% of Nepal's land area containing suitable habitat for rhesus macaques<ref name=":2">{{Cite journal |last1=Koirala |first1=S. |last2=Baral |first2=S. |last3=Garber |first3=P. A. |last4=Basnet |first4=H.|last5=Katuwal |first5=H. B. |last6=Gurung |first6=S. |last7=Rai |first7=D.|last8=Gaire |first8=R. |last9=Sharma |first9=B. |last10=Pun |first10=T. |last11=Li |first11=M. |date=2022 |title=Identifying the environmental and anthropogenic causes, distribution, and intensity of human rhesus macaque conflict in Nepal . |journal=Journal of Environmental Management|volume=316 |article-number=115276 |doi=10.1016/j.jenvman.2022.115276 |pmid=35576709 |bibcode=2022JEnvM.31615276K |s2cid=248790322}}</ref> but only having 8% of such suitable area being protected national parks.<ref name=":2" /> Rhesus macaques are rated as one of the top ten crop-raiding wildlife species in Nepal.<ref name=":2" />

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<ref name=":2" />". Mitigation strategies offers the most effective solutions to reduce conflict occurring between rhesus macaques and humans in Nepal.<ref name=":2" />

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<ref name=":6">{{Cite journal |last1=Saraswat |first1=R. |last2=Sinha |first2=Anindya |last3=R. |first3=Sindhu |date=2015 |title=A god becomes a pest? Human-rhesus macaque interactions in Himachal Pradesh, northern India |journal=European Journal of Wildlife Research |volume=61 |issue=3 |pages=435–443 |doi=10.1007/s10344-015-0913-9 |bibcode=2015EJWR...61..435S |s2cid=254194027}}</ref> with such conflict being a source of contentious debate in political scenarios, resentment and polarization amongst agriculturalists and wildlife conservationists.<ref name=":5" /> In India, crop raiding by rhesus macaques has been identified as the main cause of conflict.<ref name=":5" /> In urban areas, rhesus macaques damage property and injure people in house raids to access food and provisions;<ref name=":6" /> in agricultural areas, they cause financial losses to farmers due to crop depredation.<ref name=":6" /> The estimated extent of crop damages in Himachal Pradesh ranges from 10–100% to 40–80% of all crop losses.<ref name=":6" /> The financial implications of such damage is estimated at approximately USD$200,000 in agriculture and USD$150,000 in horticulture.<ref name=":6" /> Quantification of crop and financial loses is challenging. Farmers' negative views of macaques may cause them to perceive higher than actual losses. This has led to harsh actions against rhesus macaque communities. Other factors in rhesus perception include economic status, farmer economic stability, cultural attitudes towards the given species and the frequency and intensity of wildlife conflicts.<ref name=":6" /> All of the above have resulted in changes in conservation and management with legal rhesus macaque culling issued in 2010.<ref name=":6" />

Human-macaque conflict is also occurring in China, specifically in the area of Longyang District, Baoshan City, Yunnan Province. The peak period of conflict occurs from August–October. Factors associated with accessibility and availability of food and shelter appear to be the key drivers of human-macaque conflict, with an overall increase between the years of 2012 and 2021.<ref name=":4"/>

One key factor of conflict that directly affects 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.<ref>{{Cite journal |last1=Anand |first1=S. |last2=Vaidyanathan |first2=S. |last3=Radhakrishna |first3=S. |date=2021 |title=The Role of Landscape Structure in Primate Crop Feeding: Insights from Rhesus Macaques (''Macaca mulatta'') in Northern India |journal=International Journal of Primatology |volume=42 |issue=5 |pages=764–780 |doi=10.1007/s10764-021-00238-y |s2cid=238680232}}</ref>

Towards the end of March 2018, it was reported that a monkey had entered a house in the village of Talabasta, [[Odisha]], India 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.<ref name=BBC2018>{{cite news |work=[[BBC]] |title=Indian police search for monkey that snatched baby |url=https://www.bbc.com/news/world-asia-india-43615812 |date=2018 |access-date=2018-04-03}}</ref><ref name=9news2018>{{cite news |work=[[9news.com.au]] |title=Baby stolen by monkey in India found dead |url=https://www.9news.com.au/world/2018/04/03/05/13/baby-stolen-by-monkey-in-india-found-dead |date=2018 |access-date=2018-04-03}}</ref>

=== Population management tools ===
Crop-raiding is seen as one of the most important behaviours to change to reduce conflicts. One example is the implementation of guards in agricultural settings to chase off intruding monkeys using dogs, slingshots, and firecrackers. This method is non-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 habitats. In urban settings, planting food trees within city periphery and country parks aim to discourage macaques from entering nearby residential areas for food.<ref name=Priston2012/>
In areas of tourism, human behaviour change is necessary to prevent conflict. An important aspect is enforcing no feed regulations that only allow provisioning by trained staff at scheduled times. Regulating visitor behaviours that provoke aggressive responses from macaques, including noise regulation, greatly benefits conflict reduction.<ref name=Priston2012/>
[[Translocation (Wildlife conservation)|translocation]] of problem macaques in urban rhesus communities in India has been employed as a non-lethal solution to human–macaque conflicts.<ref name=Priston2012/> Fertility control is also feasible for reducing human–macaque conflict because it avoids killing costs and problems associated with translocation.<ref name=Priston2012/>

== In science ==
[[File:Sam prior to Little Joe 2 - C-1959-52201.jpg|thumb|[[Project Mercury]] rocket [[Little Joe 1B]], launched in 1960, carried Miss Sam to {{convert|9.3|mi|km|abbr=on}} in altitude.]]
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]], one of the elements of a person's [[Blood type|blood group]], by the discoverers of the factor, [[Karl Landsteiner]] and [[Alexander Wiener]]. The rhesus macaque was also used in the well-known experiments on maternal deprivation carried out in the 1950s by [[bioethics|controversial]] [[comparative psychology|comparative psychologist]] [[Harry Harlow]]. Other medical breakthroughs facilitated by the use of the rhesus macaque include:<ref name="Mitruka 1976">{{cite book |title=Animals for medical research: models for the study of human disease |editor1=Mitruka, B. M. |editor2=Rawnsley, H. M. |editor3=Vadehra, D. V. |publisher=Wiley & Sons |location=New York |isbn=0-89874-156-4 |author=Mitruka, B. M. |year=1976 |chapter=Introduction |pages=1–21}}</ref>
* development of the [[rabies]], [[smallpox]], 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]] launched [[monkeys and apes in space|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 program|Bion missions]]. [[Albert II (monkey)|Albert II]] became the first primate and first mammal in space during a U.S. [[V-2 rocket]] suborbital flight on 14 June 1949, and died on impact when a parachute failed.

Another rhesus monkey, Able, was launched [[Monkeys in space|on a suborbital spaceflight]] in 1959, and was among the first living beings (along with [[Miss Baker]], a [[squirrel monkey]] on the same mission) to travel in space and return alive.<ref name="Able">{{cite web |url=http://www.kansas.com/2010/03/22/1236075/kansan-among-first-to-go-to-space.html |title=Kansan among first to go to space |publisher=Wichita Eagle and Kansas.com |date=22 March 2010}}</ref>

On 25 October 1999, the rhesus macaque became the first [[cloning|cloned]] [[primate]] with the birth of [[Tetra (monkey)|Tetra]]. January 2001 had the birth of [[ANDi]], the first [[transgenic]] primate; ANDi carries foreign [[gene]]s originally from a [[jellyfish]].<ref name="GM monkey">{{cite news |title=GM monkey first |url=https://news.bbc.co.uk/2/hi/science/nature/1112171.stm |access-date=13 February 2015 |work=BBC News |agency=BBC |date=11 January 2001}}</ref>

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]] on the island of [[Cayo Santiago]], off [[Puerto Rico]], where approximately 1800 of the monkeys live.<ref>{{cite news | last =Daniel | first =Ariel | title =These monkeys were 'notoriously competitive' until Hurricane Maria wrecked their home | newspaper =[[National Public Radio]] | location = | pages = | language = | publisher = | date =July 10, 2024| url =https://www.npr.org/2024/07/09/g-s1-6734/rhesus-macaques-hurricane-maria-changed-monkey-society | access-date =November 29, 2024}}</ref> 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]]. This virus does not typically harm the monkey, but is very dangerous to humans in the rare event that it [[zoonosis|jumps species]], for example in the 1997 death of [[Yerkes National Primate Research Center]] researcher Elizabeth Griffin.<ref>{{cite web |archive-url=https://web.archive.org/web/20121101233505/http://www.ergriffinresearch.org/about-elizabeth/ |archive-date=1 November 2012 |url=http://www.ergriffinresearch.org/about-elizabeth/ |title=About Elizabeth (Beth) R. Griffin (1975–1997) |publisher=Elizabeth R. Griffin Research Foundation}}</ref><ref>{{cite news |last=Bragg |first=R. |date=1997-12-14 |url=https://www.nytimes.com/1997/12/14/us/a-drop-of-virus-from-a-monkey-kills-a-researcher-in-6-weeks.html |title=A drop of virus from a monkey kills a researcher in 6 weeks |newspaper=New York Times}}</ref><ref>{{cite web |url=http://www.emory.edu/EMORY_REPORT/erarchive/1998/January/erjanuary.12/1_12_98Yerkes.html |title=Yerkes 'family' pulled together after death of young researcher from rare Herpes B infection |publisher=Emory.edu |date=1998-01-12 |access-date=2015-03-09}}</ref>

[[File:Rhesus macaque - 20160710.webm|thumb|Rhesus macaque in a [[Japan]]ese zoo, 2016]]

=== Genome sequencing ===
{{Infobox genome | taxId = 215 | ploidy = diploid | chromosomes = 21 pairs | size = 3,097.37 Mb | year = 2007}}

Work on the [[genome]] of the rhesus macaque was completed in 2007, making the species the second nonhuman [[primate]] whose genome was sequenced.<ref>{{cite journal |title=A barrel of monkey genes |author1=Zahn, L. M. |author2=Jasny, B. R. |author3=Culotta, E. |author4=Pennisi, E. |author4-link=Elizabeth Pennisi |name-list-style=amp |journal=Science |date=2007 |doi=10.1126/science.316.5822.215 |volume=316 |issue=5822 |page=215 |doi-access=free}}</ref> Humans and macaques apparently share about 93% of their [[DNA sequence]] and shared a [[common ancestor]] roughly 25 million years ago.<ref>{{cite press release |title=DNA sequence of rhesus macaque has evolutionary, medical implications |publisher=Human Genome Sequencing Center |date=13 April 2007 |url=http://www.bcm.edu/news/item.cfm?newsID=853 |access-date=15 April 2007}}</ref> The rhesus macaque has 21 pairs of chromosomes.<ref>{{cite journal |title=Banding patterns of the chromosomes of the rhesus monkey (''Macaca mulatta'') |author1=Perticone, P. |author2=Rizzoni, M. |author3=Palitti, F. |author4=Di Chiara, P. |journal=Journal of Human Evolution |date=1974 |doi=10.1016/0047-2484(74)90023-2 |volume=3 |issue=4 |pages=291–295|bibcode=1974JHumE...3..291P }}</ref>

Comparison of rhesus macaques, [[Common chimpanzee|chimpanzee]]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]], University of Chicago.<ref>{{cite journal |doi=10.1126/science.1139477 |pmid=17431168 |title=Human-specific changes of genome structure detected by genomic triangulation |journal=Science |volume=316 |issue=5822 |pages=235–7 |year=2007 |last1=Harris |first1=R. A. |last2=Rogers |first2=J. |last3=Milosavljevic |first3=A. |bibcode=2007Sci...316..235H |doi-access=free}}</ref>

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.<ref>{{cite journal |pmid=17431167 |year=2007 |author1=Rhesus macaque genome sequencing analysis consortium |title=Evolutionary and biomedical insights from the rhesus macaque genome |journal=Science |volume=316 |issue=5822 |pages=222–234 |last2=Gibbs |first2=R. A. |last3=Rogers |first3=J. |display-authors=3 |last4=Katze |first4=M. G. |last5=Bumgarner |first5=R. |last6=Weinstock |first6=G. M. |last7=Mardis |first7=E. R. |last8=Remington |first8=K. A. |last9=Strausberg |first9=R. L. |last10=Venter |first10=J. C. |last11=Wilson |first11=R. K. |last12=Batzer |first12=M. A. |last13=Bustamante |first13=C. D. |last14=Eichler |first14=E. E. |last15=Hahn |first15=M. W. |last16=Hardison |first16=R. C. |last17=Makova |first17=K. D. |last18=Miller |first18=W. |last19=Milosavljevic |first19=A. |last20=Palermo |first20=R. E. |last21=Siepel |first21=A. |last22=Sikela |first22=J. M. |last23=Attaway |first23=T. |last24=Bell |first24=S. |last25=Bernard |first25=K. E. |last26=Buhay |first26=C. J. |last27=Chandrabose |first27=M. N. |last28=Dao |first28=M. |last29=Davis |first29=C. |last30=Delehaunty |first30=K. D.  |doi=10.1126/science.1139247 |bibcode=2007Sci...316..222. |doi-access=free}}</ref>

DNA microarrays are used in macaque research. For example, [[Michael Katze]] 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.<ref>{{cite journal |last=Cilloniz |first=C. |author2=Shinya, K. P. |author3=Korth, X. |author4=Proll, M. J. |author5=Aicher, S. C. |author6=Carter, L. D. |author7=Chang, V. S. |author8=Kobasa, J. H. |author9=Feldmann, D. |author10=Strong, F. |author11=Feldmann, J. E. |author12=Kawaoka, H. |author13=Katze, Y. |author14=Michael, G. |title=Lethal influenza virus infection in macaques is associated with early dysregulation of inflammatory related genes |journal=PLOS Pathogens |year=2009 |volume=5 |issue=10 |article-number=e1000604 |doi=10.1371/journal.ppat.1000604 |pmid=19798428 |pmc=2745659 |doi-access=free }}</ref><ref>{{cite journal |last=Brown |first=J. N. |author2=Palermo, R. E. B. |author3=Gritsenko, C. R. |author4=Sabourin, M. |author5=Long, P. J. |author6=Sabourin, J. P. |author7=Bielefeldt-Ohmann, C. L. |author8=García-Sastre, H. |author9=Albrecht, A. |author10=Tumpey, R. |author11=Jacobs, T. M. |author12=Smith, J. M. |author13=Katze, R. D. |author14=Michael, G. |title=Macaque proteome response to highly pathogenic avian influenza and 1918 reassortant influenza virus infections |journal=Journal of Virology |date=2010 |volume=84 |issue=22 |pages=12058–12068 |doi=10.1128/jvi.01129-10 |pmid=20844032 |pmc=2977874}}</ref>

The full sequence and annotation of the macaque genome is available on the [[Ensembl]] genome browser.<ref>{{Cite web |url=http://www.ensembl.org/Macaca_mulatta/Info/Index/ |title=''Macaca mulatta'' - Ensembl genome browser 96}}</ref>

== Conservation status ==
The rhesus macaque is listed as [[Least Concern]] on the [[IUCN Red List]] and estimated to exist in large numbers; it is tolerant of a broad range of [[habitat]]s, including urban environments.<ref name=iucn/> It has the largest natural range of any non-human primate.<ref name=Cooper2022/> 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.<ref>{{cite journal |last1=Malaivijitnond |first1=S. |last2=Takenaka |first2=O. |last3=Kawamoto |first3=Y. |last4=Urasopon |first4=N. |last5=Hadi |first5=I. |last6=Hamada |first6=Y. |year=2007 |title=Anthropogenic macaque hybridization and genetic pollution of a threatened population |journal=Tropical Natural History |volume=7 |issue=1 |pages=11–23 |doi=10.58837/tnh.7.1.102915 |doi-access=free }}</ref><ref>{{cite journal |title=Previously unreported population of rhesus macaques ''Macaca mulatta'' in Chiang Rai province, Thailand: preliminary observations |first1=P. |last1=Kyes |first2=P. |last2=Thamsenanupap |first3=T. |last3=Tanee |first4=A. |last4=Intralawan |first5=R. C. |last5=Kyes |journal=Asian Primates Journal |year=2018 |volume=7 |issue=1 |pages=6–13 |pmc=6914309 |pmid=31844847}}</ref> Despite the wealth of information on its ecology and behaviour, little attention has been paid to its demography or population status,<ref>{{Cite journal |last1=Kumar |first1=R. |last2=Sinha |first2=A. |last3=Radhakrishna |first3=S. |date=2013 |title=Comparative Demography of Two Commensal Macaques in India: Implications for Population Status and Conservation |url=https://brill.com/view/journals/ijfp/84/6/article-p384_4.xml |journal=Folia Primatologica |volume=84 |issue=6 |pages=384–393 |doi=10.1159/000351935 |pmid=24022675 |s2cid=45152033|url-access=subscription }}</ref> which can pose a risk for future Rhesus macaque populations. The extension of its distributional limits by approximately {{cvt|3500|km2}} in southeast India caused  population stress on other species. This range extension has been caused by human intervention tactics whereby village translocation occurs from urban conflict ridden areas.<ref name=":1">{{Cite journal |last1=Radhakrishna |first1=Sindhu |last2=Sinha |first2=A. |date=2011 |title=Less than wild? Commensal primates and wildlife conservation |journal=Journal of Biosciences |volume=36 |issue=5 |pages=749–753 |doi=10.1007/s12038-011-9145-7 |pmid=22116271 |s2cid=32085979}}</ref>

==See also==
* [[Girneys]]

== References ==
{{Reflist}}

== External links ==
{{Wikispecies|Macaca mulatta|the rhesus macaque}}{{Commons and category|Macaca mulatta|Macaca mulatta}}
* [https://web.archive.org/web/20060507095619/http://www.arkive.org/species/GES/mammals/Macaca_mulatta/ ARKive: images and movies of the rhesus macaque ''Macaca mulatta'']
* [http://brainmaps.org Brain maps and brain atlases of rhesus macaque]
* [https://web.archive.org/web/20051230210410/http://pin.primate.wisc.edu/factsheets/entry/rhesus_macaque Primate Info Net: ''Macaca mulatta'' Factsheet]
* [http://animaldiversity.ummz.umich.edu/site/accounts/information/Macaca_mulatta.html University of Michigan Museum of Zoology's Animal Diversity Web: ''Macaca mulatta'']
* {{usurped|1=[https://web.archive.org/web/20190118170107/http://www.rhesusgenome.org/ ''Macaca mulatta'' Genome]}}
* [http://www.media-generation.com/DVD%20PAGES/Rhesus/Rhesus.htm Rhesus Play] Film analysis of agonistic play by Donald Symons (UCSB) on DVD
* View the [http://www.ensembl.org/Macaca_mulatta/Info/Index/ Macaque genome] in [[Ensembl]].
* {{UCSC genomes|rheMac8}}

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