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"Macaca" redirects here. For a racial slur for dark-skinned people, see Macaca (term).

Macaques[1]
Bonnet macaque in Manegaon, Maharashtra, India
Scientific classification Edit this classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Primates
Suborder: Haplorhini
Infraorder: Simiiformes
Family: Cercopithecidae
Subfamily: Cercopithecinae
Tribe: Papionini
Genus: Macaca
Lacépède, 1799
Type species
Simia inuus[1]
Linnaeus, 1766
(= Simia sylvanus Linnaeus, 1758)
Species

See text

The macaques (/məˈkɑːk, -ˈkæk/)[2] constitute a genus (Macaca) of gregarious Old World monkeys of the subfamily Cercopithecinae. The 23 species of macaques inhabit ranges throughout Asia, North Africa, and Europe (in Gibraltar). Macaques are principally frugivorous (preferring fruit), although their diet also includes seeds, leaves, flowers, and tree bark. Some species such as the long-tailed macaque (M. fascicularis; also called the crab-eating macaque) will supplement their diets with small amounts of meat from shellfish, insects, and small mammals. On average, a southern pig-tailed macaque (M. nemestrina) in Malaysia eats about 70 large rats each year.[3][4] All macaque social groups are arranged around dominant matriarchs.[5]

Macaques are found in a variety of habitats throughout the Asian continent and are highly adaptable. Certain species are synanthropic, having learned to live alongside humans, but they have become problematic in urban areas in Southeast Asia and are not suitable to live with, as they can carry transmittable diseases.

Most macaque species are listed as vulnerable to critically endangered on the International Union of the Conservation of Nature (IUCN) Red List.

Description

[edit]

Aside from humans (genus Homo), the macaques are the most widespread primate genus, ranging from Japan to the Indian subcontinent, and in the case of the Barbary macaque (Macaca sylvanus), to North Africa and Southern Europe. Twenty-three macaque species are currently recognized. Macaques are robust primates whose arms and legs are about the same in length. The fur of these animals is typically varying shades of brown or black and their muzzles are rounded in profile with nostrils on the upper surface. The tail varies among each species, which can be long, moderate, short or totally absent.[6] Although several species lack tails, and their common names refer to them as apes, these are true monkeys, with no greater relationship to the true apes than any other Old World monkeys. Instead, this comes from an earlier definition of 'ape' that included primates generally.[7]

In some species, skin folds join the second through fifth toes, almost reaching the first metatarsal joint.[8] The monkey's size differs depending on sex and species. Males from all species can range from 41 to 70 cm (16 to 27.5 in) in head and body length, and in weight from 5.5 to 18 kg (12 to 40 lb).[6] Females can range from a weight of 2.4 to 13 kg (5.3 to 28.7 lb). These primates live in troops that vary in size, where males dominate, however the order of dominance frequently shifts. Female dominance lasts longer and depends upon their genealogical position. Macaques are able to swim and spend most of their time on the ground and spend some time in trees. They have large pouches in their cheeks where they carry extra food. They are considered highly intelligent and are often used in the medical field for experimentation due to their remarkable similarity to humans in emotional and cognitive development. Extensive experimentation has led to the long-tailed macaque being listed as endangered.[6]

Distribution and habitat

[edit]

Macaques are highly adaptable to different habitats and climates and can tolerate a wide fluctuation of temperatures and live in varying landscape settings. They easily adapt to human-built environments and can survive well in urban settings if they are able to obtain food. They can also survive in completely natural settings absent of humans.

The ecological and geographic ranges of the macaque are the widest of any non-human primate. Their habitats include the tropical rainforests of Southeast Asia, Sri Lanka, India, arid mountains of Pakistan and Afghanistan, and temperate mountains in Algeria, Japan, China, Morocco, and Nepal. Some species also inhabit villages and towns in cities in Asia.[9] There is also an introduced population of rhesus macaques in the US state of Florida consisting, essentially, of monkeys abandoned when a failed boat ride-safari was shut down in the mid-20th century.

A probable Early Pliocene macaque molar from the Red Crag Formation (Waldringfield, United Kingdom), represents one of the oldest and northernmost records of the genus in Europe reported to date.[10]

Ecology and behavior

[edit]

Diet

[edit]

Macaques are mainly frugivorous, although some species have been observed feeding on insects. In natural habitats, they have been observed to consume certain parts of over one hundred species of plants including the buds, fruit, young leaves, bark, roots, and flowers. When macaques live amongst people, they raid agricultural crops such as wheat, rice, or sugarcane; and garden crops like tomatoes, bananas, melons, mangos, or papayas.[11] In human settings, they also rely heavily on direct handouts from people. This includes peanuts, rice, legumes, or even prepared food.

Group structure

[edit]

Macaques live in established social groups that can range from a few individuals to several hundred, as they are social animals. A typical social group possess between 20 and 50 individuals of all ages and of both sexes. The typical composition consists of 15% adult males, 35% adult females, 20% infants, and 30% juveniles, though there exists variation in structure and size of groups across populations.[citation needed]

The premotor cortex of macaques is widely studied.[12]

Macaques have a very intricate social structure and hierarchy, with different classifications of despotism depending on species.[13] If a macaque of a lower level in the social chain has eaten berries and none are left for a higher-ranking macaque, then the one higher in status can, within this social organization, remove the berries from the other monkey's mouth.[14]

Reproduction and mortality

[edit]

The reproductive potential of each species differs. Populations of the rhesus macaque can grow at rates of 10% to 15% per year if the environmental conditions are favorable. However, some forest-dwelling species are endangered with much lower reproductive rates.[citation needed] After one year of age, macaques move from being dependent on their mother during infancy, to the juvenile stage, where they begin to associate more with other juveniles through rough tumble and playing activities. They sexually mature between three and five years of age. Females will usually stay with the social group in which they were born; however, young adult males tend to disperse and attempt to enter other social groups. Not all males succeed in joining other groups and may become solitary, attempting to join other social groups for many years.[citation needed] Macaques have a typical lifespan of 20 to 30 years.

As invasive species

[edit]
M. fascicularis on a scooter at Ko Chang, Thailand

Certain species under the genus Macaca have been transplanted to different parts of the world, where they become invasive, while others remain threatened. The long-tailed macaque (M. fascicularis) is listed as an invasive alien species in Mauritius, along with the rhesus macaques (M. mulatta) in Florida.[15] Despite this, the former is listed as endangered.

The long-tailed macaque causes severe damage to parts of its range where it has been introduced because the populations grow unchecked due to a lack of predators.[16] On the island of Mauritius, they have created serious conservation concerns for other endemic species. They consume seeds of native plants and aid in the spread of exotic weeds throughout the forests. This changes the composition of the habitats and allows them to be rapidly overrun by invasive plants.

Long-tailed macaques are also responsible for the near extinction of several bird species on Mauritius by destroying the nests of the birds and eating the eggs of critically endangered species, such as the pink pigeon and Mauritian green parrot.[17] They have been known to eat crops from human farms and gardens, sparking human-wildlife conflicts.

In Florida, a group of rhesus macaques inhabit Silver Springs State Park. Humans often feed them, which may alter their movement and keep them close to the river on weekends where high human traffic is present.[15] The monkeys can become aggressive toward humans, largely due to human ignorance of macaque behavior. They carry potentially fatal human diseases, including the herpes B virus.[18]

Relations with humans

[edit]

Several species of macaque are used extensively in animal testing, particularly in the neuroscience of visual perception and the visual system.

Nearly all (73–100%) captive rhesus macaques are carriers of the herpes B virus. This virus is harmless to macaques, but infections of humans, while rare, are potentially fatal, a risk that makes macaques unsuitable as pets.[19]

Urban performing macaques also carried simian foamy virus, suggesting they could be involved in the species-to-species jump of similar retroviruses to humans.[20]

Population control

[edit]

Management techniques have historically been controversial, and public disapproval can hinder control efforts. Previously, efforts to remove macaque individuals were met with public resistance.[15] One management strategy that is currently being explored is that of sterilization. Natural resource managers are being educated by scientific studies in the proposed strategy. Effectiveness of this strategy is estimated to succeed in keeping populations in check. For example, if 80% of females are sterilized every five years, or 50% every two years, it could effectively reduce the population.[15] Other control strategies include planting specific trees to provide protection to native birds from macaque predation, live trapping, and the vaccine porcine zona pellucida (PZP), which causes infertility in females.[17]

Cloning

[edit]
See also: List of animals that have been cloned

In January 2018, scientists in China reported in the journal Cell the first creation of two crab-eating macaque clones, named Zhong Zhong and Hua Hua, using somatic cell nuclear transfer – the same method that produced Dolly the sheep.[21][22][23][24]

Species

[edit]
Genus Macaca – Lacépède, 1799 – 24 species
Common name Scientific name and subspecies Range Size and ecology IUCN status and estimated population
Arunachal macaque

Brown monkey

M. munzala
Sinha, Datta, Madhusudan, Mishra, 2005 		Eastern Himalayas
Map of range 		Size: 36–77 cm (14–30 in) long, plus about 9–20 cm (4–8 in) tail[25]

Habitat: Forest[26]

Diet: Fruit, leaves, grains, buds, seeds, flowers, and bark, as well as insects and small invertebrates[25] 		 EN 


250 Population declining[26]

Assam macaque

Brown monkeys

M. assamensis
(McClelland, 1840)

Two subspecies
  • M. a. assamensis (Eastern Assamese macaque)
  • M. a. pelops (Western Assamese macaque)
Southeastern Asia
Map of range 		Size: 36–77 cm (14–30 in) long, plus about 9–20 cm (4–8 in) tail[25]

Habitat: Forest[27]

Diet: Fruit, leaves, grains, buds, seeds, flowers, and bark, as well as insects and small invertebrates[25] 		 NT 


Unknown Population declining[27]

Barbary macaque

Brown monkey and baby

M. sylvanus
(Linnaeus, 1758) 		Northwestern Africa
Map of range 		Size: 45–60 cm (18–24 in) long, plus 1–2 cm (0–1 in) tail[28]

Habitat: Forest, shrubland, grassland, rocky areas, and caves[29]

Diet: Plants, caterpillars, fruit, seeds, roots, and fungi[28] 		 EN 


Unknown Population declining[29]

Bonnet macaque

Brown monkey

M. radiata
(Geoffroy, 1812)

Two subspecies
  • M. r. diluta (Pale-bellied bonnet macaque)
  • M. r. radiata (Dark-bellied bonnet macaque)
Southern India
Map of range 		Size: 36–77 cm (14–30 in) long, plus about 9–20 cm (4–8 in) tail[25]

Habitat: Forest, savanna, and shrubland[30]

Diet: Fruit, foliage, and insects, as well as bird eggs and lizards[31] 		 VU 


Unknown Population declining[30]

Booted macaque

Drawing of gray monkey

M. ochreata
(Ogilby, 1841) 		Island of Sulawesi in Indonesia
Map of range 		Size: 36–77 cm (14–30 in) long, plus about 1–15 cm (0–6 in) tail[25]

Habitat: Forest and savanna[32]

Diet: Fruit, leaves, grains, buds, seeds, flowers, and bark, as well as insects and small invertebrates[25] 		 VU 


Unknown Population declining[32]

Celebes crested macaque

Gray monkey

M. nigra
(Desmarest, 1822) 		Island of Sulawesi
Map of range 		Size: 44–57 cm (17–22 in) long, plus about 2 cm (1 in) tail[33]

Habitat: Forest[34]

Diet: Fruit, as well as insects, shoots, leaves, and stems[33] 		 CR 


Unknown Population declining[34]

Crab-eating macaque

Brown monkey

M. fascicularis
(Raffles, 1821)

Ten subspecies
  • M. f. atriceps (Dark-crowned long-tailed macaque)
  • M. f. aureus (Burmese long-tailed macaque)
  • M. f. condorensis (Con Song long-tailed macaque)
  • M. f. fascicularis (Common long-tailed macaque)
  • M. f. fusca (Simeulue long-tailed macaque)
  • M. f. karimondjawae (Kemujan long-tailed macaque)
  • M. f. lasiae (Lasia long-tailed macaque)
  • M. f. philippensis (Philippine long-tailed macaque)
  • M. f. tua (Maratua long-tailed macaque)
  • M. f. umbrosus (Nicobar long-tailed macaque)
Southeastern Asia
Map of range 		Size: 40–47 cm (16–19 in) long, plus 50–60 cm (20–24 in) tail

Habitat: Forest, intertidal marine, caves, inland wetlands, grassland, shrubland, and savanna[35]

Diet: Fruit, crabs, flowers, insects, leaves, fungi, grasses, and clay[36] 		 EN 


Unknown Population declining[35]

Formosan rock macaque

Gray monkeys

M. cyclopis
Swinhoe, 1862 		Taiwan
Map of range 		Size: 36–45 cm (14–18 in) long, plus 26–46 cm (10–18 in) tail[37]

Habitat: Forest[38]

Diet: Fruit, leaves, berries, seeds, insects, and small vertebrates, buds, and shoots[37] 		 LC 


Unknown Population steady[38]

Gorontalo macaque

Black-and-white drawing of monkey

M. nigrescens
(Temminck, 1849) 		Island of Sulawesi
Map of range 		Size: 36–77 cm (14–30 in) long, plus about 1–15 cm (0–6 in) tail[25]

Habitat: Forest[39]

Diet: Fruit, leaves, grains, buds, seeds, flowers, and bark, as well as insects and small invertebrates[25] 		 VU 


Unknown Population declining[39]

Heck's macaque

M. hecki
(Matschie, 1901) 		Island of Sulawesi
Map of range 		Size: 36–77 cm (14–30 in) long, plus about 1–15 cm (0–6 in) tail[25]

Habitat: Forest and grassland[40]

Diet: Fruit, leaves, grains, buds, seeds, flowers, and bark, as well as insects and small invertebrates[25] 		 VU 


100,000 Population declining[40]

Japanese macaque

Brown monkeys

M. fuscata
(Blyth, 1875)

Two subspecies
 Japan
Map of range 		Size: 36–77 cm (14–30 in) long, plus about 1–15 cm (0–6 in) tail[25]

Habitat: Forest[41]

Diet: Fruit, seeds, flowers, nectar, leaves, and fungi[42] 		 LC 


Unknown Population steady[41]

Lion-tailed macaque

Gray and brown monkey

M. silenus
(Linnaeus, 1758) 		Southwestern India
Map of range 		Size: 40–61 cm (16–24 in) long, plus 24–38 cm (9–15 in) tail[43]

Habitat: Forest[44]

Diet: Fruit, as well as leaves, stems, flowers, buds, fungi, insects, lizards, tree frogs, and small mammals[43] 		 EN 


2,400–2,500 Population declining[44]

Moor macaque

Gray monkeys

M. maura
(Schinz, 1825) 		Island of Sulawesi
Map of range 		Size: 36–77 cm (14–30 in) long, plus about 1–15 cm (0–6 in) tail[25]

Habitat: Forest and grassland[45]

Diet: Fruit, leaves, grains, buds, seeds, flowers, and bark, as well as insects and small invertebrates[25] 		 EN 


Unknown Population declining[45]

Muna-Buton macaque


M. brunnescens
(Matschie, 1901) 		Island of Sulawesi in Indonesia
Map of range 		Size: 36–77 cm (14–30 in) long, plus about 1–15 cm (0–6 in) tail[25]

Habitat: Forest[46]

Diet: Fruit, leaves, grains, buds, seeds, flowers, and bark, as well as insects and small invertebrates[25] 		 VU 


Unknown Population declining[46]

Northern pig-tailed macaque

Gray monkey and baby

M. leonina
(Blyth, 1863) 		Southeastern Asia
Map of range 		Size: 36–77 cm (14–30 in) long, plus about 9–20 cm (4–8 in) tail[25]

Habitat: Forest[47]

Diet: Leaves, seeds, stems, roots, flowers, bamboo shoots, rice, gums, insects, larvae, termite eggs and spiders[47] 		 VU 


Unknown Population declining[47]

Pagai Island macaque

Brown monkey

M. pagensis
(G. S. Miller, 1903) 		Mentawai Islands in Indonesia
Map of range 		Size: 36–77 cm (14–30 in) long, plus about 9–20 cm (4–8 in) tail[25]

Habitat: Forest[48]

Diet: Fruit, leaves, grains, buds, seeds, flowers, and bark, as well as insects and small invertebrates[25] 		 CR 


2,100–3,700 Population declining[48]

Rhesus macaque

Brown monkey

M. mulatta
(Zimmermann, 1790) 		Southern and southeastern Asia
Map of range 		Size: 45–64 cm (18–25 in) long, plus 19–32 cm (7–13 in) tail[49]

Habitat: Forest, savanna, and shrubland[50]

Diet: Fish, crabs, shellfish, bird eggs, honeycombs, crayfish, crabs, spiders, plants, gums and pith[50] 		 LC 


Unknown Unknown[50]

Siberut macaque


M. siberu
Fuentes, 1995 		Siberut island in Indonesia
Map of range 		Size: 36–77 cm (14–30 in) long, plus about 9–20 cm (4–8 in) tail[25]

Habitat: Forest[51]

Diet: Fruit, as well as mushrooms, leaves, crabs, crayfish, pith, sap, shoots and flowers[51] 		 EN 


Unknown Population declining[51]

Southern pig-tailed macaque

Brown monkeys

M. nemestrina
(Linnaeus, 1766) 		Southeastern Asia
Map of range 		Size: 46–57 cm (18–22 in) long, plus 13–26 cm (5–10 in) tail[52]

Habitat: Forest and shrubland[53]

Diet: Fruit, insects, seeds, leaves, dirt, and fungus, as well as birds, termite eggs and larvae, and river crabs[52] 		 EN 


Unknown Population declining[53]

Stump-tailed macaque

Brown and red monkeys

M. arctoides
(I. Geoffroy, 1831) 		Southeastern Asia
Map of range 		Size: 48–65 cm (19–26 in) long, plus 3–7 cm (1–3 in) tail[54]

Habitat: Forest[55]

Diet: Fruit, seeds, flowers, roots, leaves, frogs, crabs, birds, and bird eggs[54] 		 VU 


Unknown Population declining[55]

Tibetan macaque

Brown monkey

M. thibetana
(A. Milne-Edwards, 1870)

Four subspecies
  • M. t. esau
  • M. t. guiahouensis
  • M. t. huangshanensis
  • M. t. thibetana
East China
Map of range 		Size: 36–77 cm (14–30 in) long, plus about 1–15 cm (0–6 in) tail[25]

Habitat: Forest and caves[56]

Diet: Fruit, as well as flowers, berries, seeds, leaves, stems, stalks, and invertebrates[56] 		 NT 


Unknown Population declining[56]

Tonkean macaque

Group of gray monkeys

M. tonkeana
(von Meyer, 1899) 		Island of Sulawesi
Map of range 		Size: 36–77 cm (14–30 in) long, plus about 1–15 cm (0–6 in) tail[25]

Habitat: Forest[57]

Diet: Fruit, leaves, grains, buds, seeds, flowers, and bark, as well as insects and small invertebrates[25] 		 VU 


Unknown Population declining[57]

Toque macaque

Brown monkey

M. sinica
(Linnaeus, 1771)

Three subspecies
  • M. s. aurifrons (Pale-fronted toque macaque)
  • M. s. opisthomelas (Highland toque macaque)
  • M. s. sinica (Common toque macaque)
Sri Lanka
Map of range 		Size: 36–53 cm (14–21 in) long, plus at least 36–53 cm (14–21 in) tail[25]

Habitat: Forest[58]

Diet: Fruit as well as tree flowers, buds, and leaves[59] 		 EN 


Unknown Population declining[58]

White-cheeked macaque

Brown monkey

M. leucogenys
Li, Zhao, Fan, 2015 		Northeastern India Size: 36–77 cm (14–30 in) long, plus about 9–20 cm (4–8 in) tail[25]

Habitat: Forest[60]

Diet: Fruit, leaves, grains, buds, seeds, flowers, and bark, as well as insects and small invertebrates[25] 		 EN 


Unknown Population declining[60]

Prehistoric (fossil) species

[edit]

See also

[edit]

References

[edit]

Sources

[edit]
[edit]
Revisions and contributorsEdit on WikipediaRead on Wikipedia

Macaque

View on Grokipedia
from Grokipedia
The macaques (genus Macaca) comprise a diverse group of 23 species of monkeys in the family Cercopithecidae and subfamily , characterized by their robust build, , and adaptability to varied environments. These medium-sized typically weigh between 3 and 18 kg, with males larger than females, and feature short to medium-length tails, pouch-like cheek pouches for , and ischial callosities for sitting. They are quadrupedal, semi-terrestrial animals capable of both arboreal and ground-based locomotion, and are known for their opportunistic omnivorous diet consisting primarily of fruits, leaves, seeds, , and small vertebrates. Macaques exhibit the broadest geographic range of any non-human genus, inhabiting regions across southern, eastern, and southeastern —from and to and the —as well as northwestern , where the (Macaca sylvanus) is the only representative outside . This distribution spans over 5 million square kilometers and includes diverse habitats such as tropical rainforests, temperate woodlands, grasslands, and even urban and agricultural areas near human settlements, demonstrating remarkable ecological plasticity. The genus originated in around 5.5 million years ago, dispersing to , with rapid occurring less than 5 million years ago, leading to four main phylogenetic groups: sylvanus, silenus, sinica, and fascicularis. Socially, macaques form large, multi-male/multi-female groups of 10 to over 100 individuals, with matrilineal structures where females remain in their natal groups and males disperse at maturity to avoid . These societies are organized around strict dominance hierarchies that influence access to food, mates, and grooming, with behaviors including formation, after conflicts, and vocalizations for communication. Species vary in traits such as pelage color (from golden to dark brown), facial markings, and tail length, which aid in species identification and adaptation. Notable species include the rhesus macaque (M. mulatta), widely used in biomedical research due to genetic similarities to humans, and the long-tailed macaque (M. fascicularis), valued for its role in ecological studies. Conservation efforts for macaques are critical, as habitat loss, hunting, and the pet trade threaten many populations; according to the , two species are critically endangered, five are endangered, eight are vulnerable, and the rest are of least concern or near threatened. Despite their adaptability, species like the macaques face localized extinctions, underscoring the need for protected areas and international regulations to sustain this ecologically and scientifically significant .

Taxonomy and evolution

Classification

Macaques belong to the Macaca in the Cercopithecinae of the family Cercopithecidae, which encompasses the monkeys distributed across and . This placement reflects their shared characteristics with other cercopithecines, such as baboons and guenons, while distinguishing them through specific morphological and ecological adaptations. The Macaca is one of the most speciose among , with 25 recognized extant species inhabiting diverse environments from tropical forests to temperate highlands. These species are systematically divided into four primary groups based on a combination of morphological features, such as cranial structure and pelage patterns, and genetic analyses including insertions: the fascicularis group, silenus group, sylvanus group, and sinica group. The fascicularis group includes species like the long-tailed macaque (Macaca fascicularis), noted for their adaptability to human-modified landscapes; the silenus group comprises Southeast Asian forms such as the (M. silenus); the sylvanus group is represented solely by the (M. sylvanus), the only macaque native to ; and the sinica group features East Asian species like the (M. thibetana) and (M. mulatta). This grouping aids in understanding phylogenetic relationships and evolutionary divergence within the genus. Key diagnostic traits of the genus Macaca include bilateral cheek pouches that allow temporary during , colorful ischial callosities on the that facilitate prolonged sitting, and non-prehensile tails varying in length from long and slender in like M. fascicularis to short or absent in others such as M. sylvanus. These features are typical of the but are particularly pronounced in macaques, supporting their omnivorous diet and terrestrial behaviors. Recent taxonomic revisions have refined this classification, including the description of the (Macaca munzala) as a new in the sinica group from northeastern in 2004 based on distinct pelage and cranial morphology, and the recognition of within M. fascicularis to account for regional variations, such as M. f. aurea in . Additional elevations, like Macaca leonina from a of M. nemestrina, highlight ongoing refinements driven by molecular data.

Phylogenetic history

The phylogenetic history of macaques traces back to early cercopithecines in Africa, with fossils like Victoriapithecus from the middle Miocene (approximately 15 million years ago) representing a basal ancestor to the subfamily Cercopithecinae, which includes the genus Macaca. This early lineage laid the groundwork for later divergences within Old World monkeys. The genus Macaca itself originated around 5.5 million years ago in northern Africa, diverging from other papionins (such as baboons in the genus Papio) during the late Miocene, with molecular clock estimates placing the initial split at approximately 7.0–6.7 million years ago. Fossil evidence supports this timeline, with early Macaca-like remains appearing in Eurasia by 5.9–5.3 million years ago, indicating rapid post-origin dispersal. Following their African origin, macaques underwent significant migrations facilitated by land bridges and favorable climatic conditions during the and Pleistocene epochs. The basal sylvanus group, represented today by the (Macaca sylvanus) in and a relict population in (), remained largely in but with fossil evidence of broader European distribution until historical times. Subsequent waves dispersed eastward into , where the genus diversified extensively; molecular data suggest the sylvanus group diverged first, followed by migrations leading to the sinica group on the Asian mainland and the silenus and fascicularis groups in around 5–4 million years ago. Recent genomic studies indicate that the fascicularis group originated from ancient hybridization between ancestors of the sinica and silenus groups. These movements were enabled by tectonic connections like the , allowing colonization of diverse environments from temperate forests to tropical islands. Key evolutionary adaptations underpinned this expansion, particularly an omnivorous diet that included fruits, seeds, leaves, insects, and opportunistic scavenging, providing resilience in variable habitats compared to more specialized folivorous or frugivorous primates. Complementing this, macaques exhibited remarkable social flexibility, with multimale-multifemale group structures that could adjust in size and composition (from 10 to over 100 individuals) to match resource availability and predation risks, facilitating survival across ecological gradients. These traits, evident in both fossil dental morphology and modern behaviors, contributed to the genus's success, resulting in 25 extant species distributed over a vast range exceeding 5 million square kilometers. The major phylogenetic branches—sylvanus (North Africa/Europe), sinica (Asian mainland, e.g., rhesus macaque), fascicularis (Southeast Asia, e.g., long-tailed macaque), and silenus (insular Southeast Asia, e.g., lion-tailed macaque)—reflect this adaptive radiation, with mitogenomic analyses confirming their monophyly and sequential divergences.

Physical characteristics

Morphology and anatomy

Macaques exhibit a quadrupedal form adapted for versatile locomotion across terrestrial and arboreal environments, supported by robust limbs with strong musculature that enable efficient walking, running, and . Their forelimbs and hindlimbs are of comparable length, facilitating a , while the skeletal structure includes a flexible spine and powerful shoulder and pelvic girdles for during movement. The hands and feet of macaques are highly prehensile, each bearing five digits with flat nails rather than claws; the thumbs and big toes are opposable, allowing precise grasping of branches, items, and substrates. This is enhanced by keratinized palms and soles featuring epidermal ridges for improved traction and sensory feedback. The overall limb configuration reflects an evolutionary balance between mobility and manipulation capabilities. Facial morphology in macaques includes a prognathic muzzle that houses the nasal and oral structures, forward-facing eyes that provide stereoscopic vision essential for in complex habitats, and a naked face with relatively hairless, pigmented capable of conveying expressive changes through color shifts and movements. The follows a dental formula of 2.1.2.3 in both the upper and lower jaws, totaling 32 teeth in adults, with bilophodont molars featuring two transverse ridges that efficiently grind fibrous and other tough material. Internally, macaques possess enlarged salivary glands, including prominent parotid and submandibular glands, which produce secretions that moisten and aid in the temporary storage of food within expandable pouches lining the buccal cavity. Macaques also feature ischial callosities, hardened pads on the that provide comfort during prolonged sitting on hard surfaces. Their vocal tract, comprising a descended , flexible , and mobile tongue, supports the generation of a diverse array of calls ranging from coos to screams. manifests primarily in body size, with males typically larger than females across macaque .

Size, coloration, and variation

Macaques exhibit considerable variation in body size across , with average head and body lengths ranging from 40 to 70 cm, excluding the tail, and weights typically between 3 and 18 kg. is pronounced, with males generally 30-50% larger than females in both length and weight, a pattern more evident in terrestrial species than arboreal ones. Fur coloration in macaques is typically grayish-brown, providing in their varied environments, though it shows species-specific diversity such as the golden pelage of lion-tailed macaques (Macaca silenus). Tail length varies markedly, from absent in the (Macaca sylvanus) to exceeding body length in species like the long-tailed macaque (Macaca fascicularis). Age and sex influence physical traits, with juveniles displaying lighter fur coloration compared to adults and less pronounced that develops post-maturity. In females, anogenital swelling occurs during estrus, serving as a visual signal of reproductive status and varying in size and intensity across . Intraspecific variation in macaque size often follows clinal patterns related to latitude, consistent with , where individuals in higher latitudes tend to be larger, as observed in Japanese macaques (Macaca fuscata) with increased body and trunk length northward. Habitat conditions can subtly influence coloration for adaptive , though this is secondary to genetic factors.

Distribution and habitat

Geographic range

Macaques (genus Macaca) are native to a broad geographic area spanning and . In , their range extends from and in the west through the , , and southern , reaching northward to and southward to the Indonesian archipelago, including numerous islands. The only native African species is the (Macaca sylvanus), which is restricted to fragmented cedar forests in the of and . This distribution makes macaques the most widespread nonhuman genus, occupying latitudes from about 40°N to 10°S. The historical range of macaques reflects an African origin for the around 7–9 million years ago, followed by dispersal into approximately 5.5 million years ago during a period of diversification. Subsequent radiations, including westward and northward expansions, occurred around 3 million years ago, leading to the establishment of multiple clades and isolated island populations through natural migrations and vicariance events associated with tectonic changes and fluctuations. Fossil evidence supports early presence in both regions, with macaques once more continuously distributed across before Pleistocene isolations. Current populations across the approximately 23 macaque species number in the tens of millions, though precise global totals are challenging due to varying survey methods; for example, the long-tailed macaque alone is estimated at around 1 million individuals. Densities are highest in the tropical forests and human-modified landscapes of and , where species like the rhesus and long-tailed macaques thrive in large groups, often exceeding 50 individuals per square kilometer in optimal areas. These populations exhibit varying degrees of fragmentation due to habitat variability within their ranges. Introduced populations have established outside the native range since the 17th–20th centuries, primarily through human activities such as pet trade, research escapes, and deliberate releases. Notable examples include rhesus macaques in , USA, originating from imports in the 1930s for biomedical purposes and now numbering several hundred in forested areas; Barbary macaques in , , descended from North African imports dating back centuries and maintained at around 230 individuals; and long-tailed macaques in , introduced by Dutch settlers around 1602, where they have become invasive with populations in the thousands. Additional feral groups exist in places like and in .

Habitat preferences

Macaques of the genus Macaca primarily inhabit tropical and subtropical forests across , , and , but the group exhibits exceptional ecological versatility, occupying a broad spectrum of niches including mangroves, coastal swamps, grasslands, semi-arid scrublands, and montane forests. This adaptability allows them to persist in both primary and secondary growth areas, often favoring disturbed habitats where human activity has altered the landscape. For example, long-tailed macaques (Macaca fascicularis) thrive in riverine zones, forest peripheries, and agricultural edges from up to 1,800 meters. A core set of environmental requirements underpins macaque selection: reliable access to freshwater sources, fruiting trees and diverse for , and dense cover—such as shrubs or tall canopies—for predator avoidance and resting. These show high tolerance for edge s and anthropogenic fringes, frequently exploiting urban-adjacent areas like parks and plantations where food resources are supplemented by human waste. Their choices are influenced by seasonal variations, including monsoons in , which affect fruit availability and prompt shifts between arboreal and terrestrial . Altitudinally, macaques range from coastal lowlands to high elevations, with rhesus macaques (Macaca mulatta) documented up to over 4,000 meters in the , while species like the (Macaca thibetana) occupy subtropical to temperate mixed forests between 800 and 2,500 meters. Climatically, they span tropical rainforests to cool temperate zones, demonstrating physiological resilience to temperature extremes. In microhabitats, forested species tend to be semi-arboreal, climbing for fruits and escape, but become predominantly ground-dwelling in open grasslands; notably, Japanese macaques (Macaca fuscata) incorporate hot springs into their winter routines for in subalpine environments up to 2,900 meters. These preferences align with resource distribution, as fruit abundance drives occupancy in fruit-rich forests.

Behavior and ecology

Social organization

Macaques typically live in matrilineal societies characterized by multi-male, multi-female troops ranging from 10 to over 200 individuals, depending on the species and ; for instance, rhesus macaques (Macaca mulatta) form groups of 20 to 100 members, while some populations of long-tailed macaques (Macaca fascicularis) can exceed 150. Females are philopatric, remaining in their natal groups and forming stable matrilines that often span multiple generations, whereas males disperse at to join other groups, reducing . This structure fosters complex social networks where plays a central role in alliances and resource access. Dominance hierarchies in macaques vary by species, ranging from despotic systems with steep, aggressive rank gradients to more egalitarian ones with tolerant interactions; rhesus and Japanese macaques (Macaca fuscata) exhibit despotic hierarchies, while Barbary macaques (Macaca sylvanus) show greater tolerance. Female hierarchies are generally linear and stable, inherited matrilineally with higher-ranking mothers conferring rank to daughters, promoting kin-biased support during conflicts. In contrast, male hierarchies are more fluid and contest-based, often involving coalitions, challenges, and rank reversals, particularly among immigrants seeking mating opportunities. Communication among macaques is multifaceted, relying on vocalizations such as coos for maintaining contact and screams during agonistic encounters, alongside facial expressions like grimaces to signal submission and lip-smacking for affiliation. Grooming serves as a primary bonding mechanism, reinforcing social ties and reducing tension, especially within matrilines, while olfactory signals, including marking, convey information about dominance, identity, and reproductive status. include fission-fusion patterns, where troops split into temporary foraging subgroups to optimize resource use before reuniting, and by incoming males, which accelerates female estrus by eliminating unrelated offspring and thereby enhances the killers' .

Diet and foraging

Macaques are opportunistic omnivores, with their diet primarily consisting of plant matter such as fruits, leaves, , flowers, and bark, which typically comprises 50-80% of their intake depending on and availability. This frugivorous foundation is supplemented by animal-derived foods, including , small vertebrates like and birds, fungi, and occasionally eggs or crabs, allowing flexibility in resource-poor environments. In areas near human settlements, macaques readily incorporate anthropogenic food waste, such as grains, , and processed items, which can constitute a significant portion of their diet in urban or agricultural settings. Foraging behaviors in macaques combine ground-level searches with arboreal exploration, enabling them to exploit diverse microhabitats within their range. Species like the long-tailed macaque (Macaca fascicularis) demonstrate advanced tool use, employing stones to crack open nuts, shellfish, or other hard-shelled prey during extractive foraging along coastal or intertidal zones. These strategies are often solitary or small-group activities, with individuals scanning for opportunistic items while minimizing energy expenditure through efficient scanning and manipulation. Macaques exhibit seasonal adaptations to fluctuations in food availability, shifting to fallback resources such as bark, mature leaves, or lichens when preferred fruits become scarce during dry or winter periods. Prominent cheek pouches facilitate this by allowing individuals to collect and transport food items to safer locations for consumption, reducing exposure to predators or competitors during extended bouts. This pouch usage is particularly vital in patchy environments, where rapid collection enhances survival during scarcity. Physiologically, macaques are fermenters with an enlarged that supports microbial breakdown of fibrous plant materials, extracting from cellulose-rich fallback foods through . Their digestive system features relatively rapid gut transit times, suited to a mixed omnivorous diet that includes easily digestible fruits alongside tougher , optimizing absorption without prolonged retention. This adaptation underscores their ecological versatility across tropical and temperate habitats.

Reproduction and development

Macaques exhibit a polygynandrous , in which both males and females mate with multiple partners during the breeding season, leading to low paternity certainty and opportunities for post-copulatory . In many species, particularly those in temperate regions like rhesus macaques (Macaca mulatta), females ovulate seasonally, with conception peaks typically occurring in winter to align births with periods of resource abundance in spring and summer. lasts 5-6 months, averaging 164 days in rhesus macaques and 167 days in cynomolgus macaques (Macaca fascicularis), and results in the birth of a single offspring, though twins occur rarely (less than 1% of cases). Newborn macaques are altricial, born with limited mobility and dependent on their mothers for ; infants cling to the mother's ventral surface immediately after birth and begin suckling within the first hour. Maternal care is intensive during the first 6-12 months, involving , grooming, and protection, with typically occurring around 12 months, though nutritional begins earlier at 4-6 months as infants start solid foods. , or care provided by non-maternal group members such as female kin or subadult siblings, is common and helps reduce maternal energetic costs while enhancing infant through additional vigilance and handling. Male involvement in is generally minimal across macaque , limited to occasional guarding against predators or in like Barbary macaques (Macaca sylvanus), without direct provisioning or carrying. Infants achieve locomotor independence by 6-8 weeks and social integration into the group by 3-6 months, but remain vulnerable, with juvenile mortality rates ranging from 20-50% in the wild due primarily to predation, disease, and accidents. is reached at 3-5 years for females and 4-7 years for males, varying by species and environmental conditions; for example, rhesus females mature at about 3 years, while males do so at 4 years. In the wild, the lifespan of macaques is less than 15 years, with fewer than 5% surviving beyond 25 years, due to cumulative risks, whereas in captivity, lifespans extend to a of over 25 years and a maximum of 40 years, reflecting reduced predation and better veterinary care.

Interactions with humans

As invasive species

Macaque species have established invasive populations outside their native ranges in several regions, including Florida in the United States, Gibraltar in Europe, and various Pacific islands such as Angaur in Palau. In Florida, rhesus macaques (Macaca mulatta) were intentionally introduced in the 1930s to an island in Silver Springs State Park by a tour boat operator to attract visitors, with initial releases of about six individuals followed by additional ones in the 1940s. These populations have since expanded, with estimates of nearly 200 to 300 individuals across five groups as of 2025, driven by the absence of natural predators and access to human-provided food sources like tourist scraps and nearby agriculture. In Gibraltar, Barbary macaques (Macaca sylvanus), introduced centuries ago likely by Moorish settlers, maintain a stable population of around 230 individuals that roam urban areas, benefiting from a lack of predators and supplemental feeding by tourists. On Pacific islands like Angaur in Palau, long-tailed macaques (Macaca fascicularis) were introduced in the early 20th century during German colonial rule and further spread via Japanese and U.S. military activities during World War II, leading to populations of about 5,000 that outnumber the island's 114 human residents by approximately 44 to 1 as of 2025 due to similar factors of no predators and reliance on human food waste. The spread of these invasive macaque populations often results from intentional releases for purposes such as biomedical research, tourism attractions, or as pets, as well as accidental escapes from facilities. In , early releases were tourism-driven, but subsequent biomedical interests led to trapping and relocation of hundreds of individuals between 1998 and 2012, inadvertently aiding dispersal to form additional groups. In the Pacific, military releases during wartime contributed to establishment on islands like , where macaques have since dispersed across the 4-square-kilometer landmass. Dispersal rates can reach several kilometers per year, with documented group movements and home ranges averaging 1-2 km daily in human-modified habitats, facilitating expansion into new areas without natural barriers. Invasive macaques exert significant ecological and economic impacts, including crop raiding that causes substantial agricultural losses, competition with native wildlife, and potential hybridization risks in regions with related species. In areas like and , raiding of fruits, vegetables, and grains leads to estimated losses of up to 20% in affected farms, exacerbating issues on small islands where monkeys consume or destroy limited resources. These compete with native birds and small mammals for fruits and seeds, potentially disrupting local , as seen with long-tailed macaques preying on eggs and displacing avian species in Pacific habitats. Hybridization poses additional threats where invasive macaques overlap with native , such as rhesus macaques interbreeding with Japanese macaques (Macaca fuscata) in introduced Asian contexts, leading to genetic dilution and reduced fitness in endemic populations. Management of invasive macaque populations involves challenges like implementing sterilization programs and , which are often controversial due to public sympathy for these charismatic animals and risks of transmission. In Florida's Silver Springs, ongoing proposals include sterilization of adult females or to manage growth, as the population has exceeded earlier predictions and continues to increase as of 2025, but face opposition amid concerns over herpes B virus, which is fatal to humans and prevalent in about 25% of the monkeys. Similarly, in , trapping and contraception efforts have failed to curb growth, with debated due to ethical issues and potential zoonotic spread, highlighting the tension between ecological control and societal perceptions. These strategies underscore the difficulty in balancing management with and conservation .

Uses in research and medicine

Macaques, particularly rhesus (Macaca mulatta) and cynomolgus (Macaca fascicularis) species, serve as key model organisms in biomedical research due to their physiological and genetic similarities to humans. Rhesus macaques have been extensively used in since the 1960s, notably in studies mapping the , where researchers like David Hubel and demonstrated columnar organization in response to visual stimuli, laying foundational insights into . These models continue to inform investigations into neural circuits, such as dendritic in . Cynomolgus macaques, meanwhile, are prominent in vaccine development, including trials for and , where they replicate human-like immune responses to assess efficacy and safety; for instance, they have been used to evaluate vaccine candidates, showing comparable infection models to rhesus macaques. In , macaques have marked significant milestones, including the first successful of a via (SCNT) in 2018, when Chinese researchers produced two long-tailed macaque (Macaca fascicularis) clones named , overcoming barriers to that had previously limited . This achievement built on earlier efforts and has advanced understanding of embryonic development and potential therapeutic techniques, though it raised questions about efficiency and viability. Ethical considerations in macaque research emphasize the 3Rs principles—replacement, reduction, and refinement—introduced by Russell and Burch in 1959 to minimize animal use and suffering, which are now integral to protocols for non-human studies. Globally, approximately 100,000 macaques are used annually in laboratories, primarily for biomedical purposes, though alternatives like human-derived organoids are emerging to model brain development and reduce reliance on live animals. In the United States, the Animal Welfare Act (AWA) of 1966, amended in 1985, mandates standards for housing, veterinary care, and psychological well-being of non-human primates, including macaques, enforced by the USDA to ensure humane treatment. In the , Directive 2010/63/EU similarly requires authorization for primate procedures, promotes the 3Rs, and sets minimum enclosure sizes for like macaques to support social housing. Research involving macaques carries risks of zoonotic transmission, such as simian foamy virus (SFV), a endemic to that has infected up to 4% of occupationally exposed humans, including lab workers, through bites or mucosal contact, though it typically causes persistent infections without evident disease in humans. Regulations under the and EU Directive 2010/63/ include measures, such as and medical surveillance, to mitigate these hazards.

Zoonotic disease risks

Risks of bacterial infections or other zoonotic diseases from brief, non-injurious contact with macaques, such as casual touching while clothed, are negligible to zero. Bacterial infections require wounds for entry, while most zoonoses, including Herpes B virus, demand direct fluid exchange—such as infected saliva contacting broken skin or mucous membranes—or inhalation of aerosols or prolonged exposure. No documented transmissions have occurred from similar casual contact, consistent with CDC guidelines. Other viruses like mpox are not linked to transmission from macaques via this route.

Cultural and economic significance

Macaques hold profound religious significance in Hinduism, where species like the rhesus macaque (Macaca mulatta) are revered as embodiments of Hanuman, the monkey god from the epic Ramayana known for his strength, devotion, and loyalty to Lord Rama. Devotees across India maintain large populations of these monkeys at Hanuman temples, providing daily feedings of fruits, grains, and sweets as acts of worship, which sustains troops numbering in the hundreds at sites like the Galtaji Temple in Rajasthan or the Jagannath Temple in Puri, where over 130 rhesus macaques reside. This reverence fosters a cultural taboo against harming them, even as rhesus macaques cause substantial crop damage in agricultural areas, leading to conflicts where farmers report losses but hesitate to use lethal controls due to religious beliefs. In East Asian folklore, macaques inspire iconic figures such as Sun Wukong, the Monkey King from the 16th-century Chinese novel , depicted as a powerful, mischievous macaque-like deity with shape-shifting abilities and a staff weapon, symbolizing rebellion, immortality, and Buddhist enlightenment. This character, born from a stone egg in a macaque troop, has permeated literature, theater, and modern media, influencing global pop culture through adaptations like (e.g., Dragon Ball's ) and films that portray intelligent, anthropomorphic . While not direct models, macaques' social intelligence and adaptability have informed primate representations in Western cinema, such as the hierarchical ape societies in the franchise, which draws on real macaque behaviors observed in studies for dramatic tension. Economically, macaques drive in several regions, boosting local economies through . In , the only wild population of Barbary macaques (Macaca sylvanus) in attracts approximately 700,000 visitors annually to the Upper Rock , where guided tours and feeding sessions generate revenue for conservation and hospitality sectors, with the macaques serving as a attraction since the . Similarly, in , , the Sacred Forest Sanctuary in , home to around 700 long-tailed macaques (Macaca fascicularis), draws approximately 120,000 visitors per year, with entry fees and surrounding businesses contributing significantly to the local economy, estimated at hundreds of thousands of dollars annually from direct alone. However, the illegal pet trade undermines these benefits, as poaching for exotic pets persists despite international bans under ; for instance, 200–300 Barbary macaques are trafficked yearly from , fueling a worth millions globally and exacerbating population declines. Historically, macaques were traded across the Mediterranean for entertainment in , where Barbary macaques were imported as exotic pets and performers in circuses and private villas, often dressed in clothing or trained for tricks to amuse elites, as evidenced by mosaics and tomb inscriptions from sites like . This trade, documented from the BCE, highlighted their status as symbols of luxury and exoticism, with some receiving elaborate burials alongside human owners, reflecting their integration into Roman society before stricter import controls emerged.

Conservation

Threats and status

Macaque species face significant conservation threats, primarily from habitat loss driven by across their native ranges in and . In , where most species occur, forest cover has declined at an average rate of 0.6% per year between 1990 and 2010, resulting in a net loss of 1.6 million hectares annually and severely fragmenting macaque habitats essential for their survival. This loss is largely attributed to , logging, and infrastructure development, affecting over 76% of threatened species including macaques. Hunting for , , and the pet trade further exacerbates population declines. In regions like , intensive hunting targets macaques for consumption and medicinal uses, contributing to local extirpations and overall range reductions. The international pet trade, often involving of infants after mothers are killed, has decimated populations of species such as the long-tailed macaque (Macaca fascicularis), with thousands seized annually in illegal shipments. In October 2025, the IUCN reaffirmed the long-tailed macaque's Endangered status following a review, citing an inferred 50–70% over the past three generations due to habitat loss and exploitation. According to the , of the 24 macaque species, two are classified as Critically Endangered (e.g., the , Macaca nigra, and the , Macaca silenus), nine as Endangered, eight as Vulnerable, and five as Near Threatened or Least Concern, with the majority showing declining population trends. Global wild macaque populations are estimated in the millions but remain fragmented, with many species persisting in small, isolated groups vulnerable to events. For instance, the (Macaca silenus) numbers fewer than 4,000 individuals, confined to remnant forest patches in India's . In contrast, adaptable species like the long-tailed macaque maintain larger populations in urban and agricultural edges, though even these are declining by up to 50% in some areas over recent generations. Emerging threats include , which is projected to alter macaque ranges by shifting suitable habitats and exacerbating resource scarcity, potentially leading to further contractions for highland species like the (Macaca thibetana). Disease outbreaks pose additional risks, with zoonotic pathogens such as capable of spilling over between macaques, bats, and humans in overlapping habitats, as demonstrated in experimental infections of cynomolgus macaques (Macaca fascicularis). While native populations decline, introduced macaque groups in non-native regions, such as parts of and beyond, are expanding as notable exceptions.

Protection and management

Macaque species receive varying levels of international protection under the Convention on International Trade in Endangered Species of Wild Fauna and Flora (), with most listed in Appendix II, requiring export permits to regulate trade and prevent overexploitation. The (Macaca silenus) is afforded stricter safeguards in Appendix I, prohibiting commercial to protect its endangered status. Similarly, the (Macaca sylvanus) was uplisted to Appendix I in 2016, implementing a global ban on commercial trade to curb poaching and trafficking from North African habitats. Nationally, protections include India's Wildlife (Protection) Act of 1972, which schedules lion-tailed macaques as a Schedule I species with the highest level of prohibition against hunting and trade, supported by reserves such as in the . In Gibraltar, the only European wild macaque population—Barbary macaques—is managed under local laws like the Animals Protection and Control Ordinance, which bans feeding to maintain natural behaviors and habitat use, though post-Brexit enforcement aligns with rather than directives. Conservation initiatives emphasize habitat restoration and conflict mitigation. Reintroduction efforts for the in Morocco's , guided by population viability analyses, aim to bolster fragmented groups through captive-bred releases and habitat enhancement, with projects like the Barbary Macaque Awareness and Conservation initiative monitoring over 100 individuals since the early to assess long-term viability. In , community-based management programs in areas like North Sulawesi's Tangkoko engage local farmers in non-lethal deterrents, such as barrier fencing and alternative crop planting, reducing long-tailed macaque (Macaca fascicularis) crop raids by up to 40% while preserving forest connectivity. Successes highlight adaptive strategies. Japan's hunting ban on Japanese macaques (Macaca fuscata) since 1947 under national wildlife laws has stabilized populations in protected areas like Jigokudani, allowing recovery from historical declines through enforced no-hunt zones and supplemental feeding controls. Genetic monitoring via non-invasive fecal sampling has proven effective for species like the (Macaca cyclopis) in , enabling accurate population estimates and kinship analysis without disturbing wild groups, informing targeted habitat corridors. Ongoing challenges include transboundary illegal trade, such as the smuggling of long-tailed macaques from into for biomedical research, complicating enforcement across borders despite quotas. Balancing with habitat integrity remains critical, as provisioning at sites like Mountain has habituated Tibetan macaques (Macaca thibetana), increasing aggression and disease transmission risks while fragmenting core foraging areas.

Species diversity

Living species

The genus Macaca includes 24 extant species of Old World monkeys, primarily native to with one species extending to and . These species are classified into seven phylogenetic groups based on molecular and morphological evidence: the M. sylvanus group (1 species), M. nemestrina group (often called silenus; 11 species), M. fascicularis group (1 species), M. arctoides group (1 species), M. mulatta group (3 species), and M. sinica group (7 species). The silenus group features species adapted to forested habitats in and , such as the (M. silenus), which has distinctive black fur, a silver-white mane, and is endemic to the of southern , where it faces and is classified as Endangered. Other silenus group members include the (M. leonina), ranging across from to in tropical forests up to 1,800 m elevation, rated Vulnerable due to hunting and ; and the (M. nemestrina), found in similar habitats from to , also Vulnerable from agricultural expansion. The sinica group encompasses seven species largely confined to the and Himalayan regions, exhibiting adaptations to diverse elevations from lowlands to high-altitude forests. For instance, the (M. thibetana), the largest macaque, inhabits temperate forests in , Bhutan, and up to 4,500 m, and is listed as Near Threatened from poaching and habitat loss. The Assamese macaque (M. assamensis) occupies evergreen forests in , southern , and , rated Endangered due to the pet and deforestation. Recent taxonomic revisions have added species like the (M. munzala), described in 2005 and restricted to high-altitude areas in , , classified as but threatened by development; the (M. leucogenys), identified in 2015 from southeastern and , Vulnerable from illegal ; and the Sela macaque (M. selai), elevated to full species status in 2022 based on , found in Arunachal Pradesh's mountainous forests and potentially at risk from isolation. Prominent species from other groups include the (M. mulatta) of the mulatta group, widespread across South and from to southern in varied s including urban areas, with body length 47–65 cm and 20–30 cm; it is rated Least Concern due to its adaptability but faces local culling as a crop pest. The long-tailed macaque (M. fascicularis) of the fascicularis group inhabits from the to the , including mangroves and coastal forests where it swims proficiently, but was uplisted to Endangered in 2022 owing to for biomedical research and conversion, with a suspected 50% over three generations. As of October 2025, the IUCN reconfirmed this Endangered status following challenges from the biomedical industry. The (M. arctoides) of the arctoides group ranges through southern , Northeast India, and in subtropical forests; it exhibits a short (5–10 cm) and brownish fur, is Vulnerable from hunting, and shows evidence of ancient hybridization with sinica and mulatta group species in contact zones, leading to . Sulawesi group species, part of the silenus , are endemic to the Indonesian island of and nearby islands, numbering eight taxa including the crested black macaque (M. nigra), with dark fur and a prominent crest, critically endangered from logging and hunting in northern Sulawesi's rainforests; and the Moor macaque (M. maura), south Sulawesi's largest , Vulnerable due to agricultural encroachment. The (M. sylvanus), the sole sylvanus group member and only wild macaque outside , lives in cedar forests of , , and at elevations up to 2,600 m, rated Endangered from degradation and tourism pressures. variation is notable, such as in the long-tailed macaque, where the Nicobar subspecies (M. f. umbrosa) is restricted to three Indian islands and classified as Vulnerable due to small population size and development threats like the Great Nicobar project. Hybridization occurs in overlap zones, as seen with stump-tailed macaques interbreeding with Assamese macaques, potentially complicating conservation genetics. Overall, most macaque species face extinction risks from habitat loss, hunting, and human-wildlife conflict, with 19 of 24 listed as Vulnerable or higher on the .

Fossil species

The fossil record of macaques (genus Macaca) provides key insights into their evolutionary history, with the earliest known species indicating an African origin prior to dispersal into . The oldest described macaque fossil is Macaca libyca, from the site of Wadi Natrun in , dated to approximately 6.2–5.0 million years ago (mya). This co-occurred with the colobine Libypithecus markgrafi, suggesting early diversification of monkeys in before the genus's radiation into during the . Several extinct macaque species highlight regional adaptations and dispersals. In Europe, Macaca majori from the Pliocene of Sardinia, Italy, represents an insular endemic form characterized by dwarfism, with dental and cranial features indicating a body size smaller than most continental macaques. This size reduction likely resulted from island dwarfing in response to limited resources on the isolated Mediterranean landmass. In Asia, Macaca anderssoni from the Early Pleistocene of Henan Province, northern China (approximately 2–1 mya), exhibits facial morphology adapted to colder climates, including a lower nasal profile that may have facilitated thermoregulation in glacial environments. Important fossil sites include the Siwalik Hills of northern and , where multiple macaque taxa are recorded from 5–2 mya, documenting the eastward migration and diversification of the during the to early . These remains, primarily dental fragments, belong to early Macaca lineages that overlapped with other cercopithecids in forested habitats. In , sites such as the Upper Valdarno Basin in have yielded Macaca florentina from around 1.5 mya, illustrating migratory pathways from via the Mediterranean during the . Evolutionary patterns inferred from the fossil record include body size reductions in insular populations, as seen in M. majori, contrasting with larger ancestral forms. Macaques became extinct in around 1 mya, likely due to cooling climates during the and increased competition from other and carnivores. This marked the retreat of the genus to refugia in and , shaping its modern distribution.

References

  1. https://www.sciencedirect.com/topics/biochemistry-genetics-and-molecular-biology/macaca
  2. https://primate.wisc.edu/primate-info-net/pin-factsheets/pin-factsheet-rhesus-macaque/
  3. https://macaques.nc3rs.org.uk/about-macaques
  4. https://pmc.ncbi.nlm.nih.gov/articles/PMC2783879/
  5. https://www.researchgate.net/publication/227924320_The_genus_Macaca_A_Review_of_taxonomy_and_evolution
  6. https://animaldiversity.org/accounts/Macaca_radiata/
  7. https://besjournals.onlinelibrary.wiley.com/doi/10.1111/1365-2656.14223
  8. https://www.sciencedirect.com/science/article/pii/B9780124171442000019
  9. https://www.sciencedirect.com/topics/agricultural-and-biological-sciences/cercopithecidae
  10. https://link.springer.com/article/10.1007/s10764-005-5333-3
  11. https://www.sciencedirect.com/science/article/pii/B9780124095274000171
  12. https://www.pnas.org/doi/10.1073/pnas.0502186102
  13. https://www.sciencedirect.com/science/article/pii/S0047248418302458
  14. https://www.science.org/doi/10.1126/sciadv.add3580
  15. https://link.springer.com/article/10.1007/BF02739365
  16. https://journals.plos.org/plosone/article?id=10.1371/journal.pone.0114099
  17. https://www.intechopen.com/chapters/78564
  18. https://samantac.agnesscott.org/uncategorized/diversity-of-life/rhesus-macaque-structure-function/
  19. https://www.science.org/doi/10.1126/sciadv.1600723
  20. https://pmc.ncbi.nlm.nih.gov/articles/PMC10092073/
  21. https://neprimateconservancy.org/rhesus-macaque/
  22. https://primate.wisc.edu/primate-info-net/pin-factsheets/pin-factsheet-long-tailed-macaque/
  23. https://barbarymacaque.blogs.lincoln.ac.uk/study-species/
  24. https://primate.wisc.edu/primate-info-net/pin-factsheets/pin-factsheet-pig-tailed-macaque/
  25. https://pmc.ncbi.nlm.nih.gov/articles/PMC3483276/
  26. https://pmc.ncbi.nlm.nih.gov/articles/PMC3335495/
  27. https://primate.wisc.edu/primate-info-net/pin-factsheets/pin-factsheet-japanese-macaque/
  28. https://www.departments.bucknell.edu/biology/resources/msw3/browse.asp?s=y&id=12100565
  29. https://genomebiology.biomedcentral.com/articles/10.1186/gb-2007-8-9-226
  30. https://macaquenet.github.io/macaque_species/
  31. https://pmc.ncbi.nlm.nih.gov/articles/PMC11122667/
  32. https://edis.ifas.ufl.edu/publication/UW412
  33. https://www.nhm.ac.uk/discover/barbary-macaques-europe-and-north-africas-last-monkeys.html
  34. https://theconversation.com/the-macaque-monkeys-of-mauritius-an-invasive-alien-species-and-a-major-export-for-research-176569
  35. https://www.bornfreeusa.org/species/macaques/
  36. https://www.iucngisd.org/gisd/species.php?sc=139
  37. https://elifesciences.org/articles/78169
  38. https://www.nature.com/articles/s41598-020-72606-2
  39. https://www.gbif.org/species/2436609
  40. https://pubmed.ncbi.nlm.nih.gov/29616368/
  41. https://www.nature.com/scitable/knowledge/library/primate-sociality-and-social-systems-58068905/
  42. http://primate.uchicago.edu/2011CayoBook02.pdf
  43. https://royalsocietypublishing.org/doi/10.1098/rstb.2024.0121
  44. https://pmc.ncbi.nlm.nih.gov/articles/PMC4898773/
  45. https://pmc.ncbi.nlm.nih.gov/articles/PMC10547472/
  46. https://royalsocietypublishing.org/doi/10.1098/rsos.250608
  47. https://journals.biologists.com/jeb/article/213/8/1338/10184/Short-term-group-fission-processes-in-macaques-a
  48. https://pmc.ncbi.nlm.nih.gov/articles/PMC9311617/
  49. https://pmc.ncbi.nlm.nih.gov/articles/PMC9532248/
  50. https://pmc.ncbi.nlm.nih.gov/articles/PMC5300017/
  51. https://animaldiversity.org/accounts/Macaca_fascicularis/
  52. https://pmc.ncbi.nlm.nih.gov/articles/PMC6072728/
  53. https://www.researchgate.net/publication/280546434_Diet_of_Long-Tailed_Macaques_Macaca_fascicularis_at_the_Entrance_of_Kuala_Selangor_Nature_Park_Anthropogenic_Habitat_Food_Selection_that_Leads_to_Human-Macaque_Conflict
  54. https://pubmed.ncbi.nlm.nih.gov/29086889/
  55. https://www.science.org/doi/10.1126/sciadv.ade8159
  56. https://www.cambridge.org/core/books/primates-in-flooded-habitats/longtailed-macaque-stone-tool-use-in-intertidal-habitats/E1FE787E7B73A67A4049F27134AB9AB2
  57. https://pmc.ncbi.nlm.nih.gov/articles/PMC10284112/
  58. https://www.ias.ac.in/article/fulltext/jbsc/048/0030
  59. https://neprimateconservancy.org/japanese-macaque/
  60. https://www.nationalgeographic.com/animals/article/cheek-pouches-stuffed-animals-news
  61. https://dukespace.lib.duke.edu/server/api/core/bitstreams/1b2c5d90-cc5e-4158-af0a-105ff97c68a8/content
  62. https://www.researchgate.net/figure/Gastro-intestinal-tract-of-the-barbary-macaque-Macaca-sylvatw-FOl-showing-a-rather_fig3_280784439
  63. https://www.researchgate.net/figure/A-Comparison-of-Gut-Passage-Times-and-Body-Weight-in-Primates_tbl1_229941994
  64. https://www.nature.com/articles/s41598-025-07399-3
  65. https://www.nidirect.gov.uk/articles/welfare-primates-need-suitable-diet
  66. https://www.sciencedirect.com/science/article/pii/S096098222030186X
  67. https://macaques.nc3rs.org.uk/about-macaques/life-history
  68. https://doi.org/10.1095/biolreprod57.2.335
  69. https://onlinelibrary.wiley.com/doi/full/10.1002/ajpa.25035
  70. https://onlinelibrary.wiley.com/doi/full/10.1002/ajp.23584
  71. https://www.clickorlando.com/health/2025/02/11/could-these-wild-florida-monkeys-give-you-herpes-heres-where-theyve-been-seen/
  72. https://springsinflorida.com/blog/why-are-there-monkeys-at-silver-springs/
  73. https://www.instagram.com/p/DEupxHEzsav/
  74. https://www.abc.net.au/pacific/programs/pacific-specials/monkeys-in-palau/105811648
  75. https://www.hawaiipublicradio.org/pacific-news-minute/2025-11-05/pacific-news-minute-monkeys-outnumber-residents-on-palau
  76. https://www.wftv.com/news/9-investigates/the-history-of-rhesus-macaques-in-marion-county/299930331/
  77. https://www.sciencedirect.com/science/article/abs/pii/S0301479721013931
  78. https://www.nies.go.jp/biodiversity/invasive/DB/detail/10390e.html
  79. https://www.nationalgeographic.com/animals/article/florida-rhesus-monkeys-herpes-running-wild-invasive-species
  80. https://wwwnc.cdc.gov/eid/article/24/2/17-1439_article
  81. https://www.mdpi.com/2079-7737/12/6/848
  82. https://www.nature.com/articles/s41467-021-23884-5
  83. https://www.nature.com/articles/s41392-021-00861-4
  84. https://www.nature.com/articles/s41541-025-01103-2
  85. https://www.nature.com/articles/d41586-024-00136-2
  86. https://pmc.ncbi.nlm.nih.gov/articles/PMC7610564/
  87. https://riseforanimals.org/news/endangered-macaques/
  88. https://www.nature.com/articles/d41586-024-02314-8
  89. https://www.ncbi.nlm.nih.gov/books/NBK232192/
  90. https://eur-lex.europa.eu/LexUriServ/LexUriServ.do?uri=OJ:L:2010:276:0033:0079:en:PDF
  91. https://pmc.ncbi.nlm.nih.gov/articles/PMC2562341/
  92. https://pmc.ncbi.nlm.nih.gov/articles/PMC353775/
  93. https://policymanual.nih.gov/3044-2
  94. https://ec.europa.eu/health/opinions/en/non-human-primates/l-3/1-use-eu.htm
  95. https://www.cdc.gov/herpes-b-virus/about/index.html
  96. https://recordsofzsi.com/index.php/zsoi/article/download/158887/109478/390820
  97. https://www.atlasobscura.com/articles/fuzz-mary-roach-excerpt-india-monkeys
  98. https://www.nationalgeographic.com/history/article/who-is-sun-wukong-monkey-king
  99. https://www.discoverwildlife.com/animal-facts/mammals/monkey-business-if-macaques-want-your-food-theyre-not-going-to-ask-for-it
  100. https://www.researchgate.net/figure/Figure-A-Each-year-700-000-tourists-visit-Gibraltars-Upper-Rock-Reserve-contributing_fig1_5254047
  101. https://villaamrita.com/monkey-forest-ubud-in-bali/
  102. https://www.sciencedirect.com/science/article/pii/S0006320725004100
  103. https://medievalmonkeys.wordpress.com/2020/07/30/animal-highlights-from-ostia-gateway-to-rome/
  104. https://www.ancient-origins.net/science-history-ancient-traditions/military-monkeys-0017402
  105. https://www.nature.com/articles/s41467-019-09646-4
  106. https://pmc.ncbi.nlm.nih.gov/articles/PMC5242557/
  107. https://wwfasia.awsassets.panda.org/downloads/primates_of_the_greater_mekong_report__2021___single_page.pdf
  108. https://news.mongabay.com/2025/10/iucn-upholds-long-tailed-macaques-endangered-status-after-complaint/
  109. https://www.bornfree.org.uk/news/mad-about-macaques-celebrating-international-macaque-week/
  110. https://mbimph.com/index.php/UPJOZ/article/download/3845/3486/6196
  111. https://www.peta.org/media/news-releases/new-studies-show-staggering-drop-in-macaque-populations-peta-pushes-government-to-act/
  112. https://www.sciencedirect.com/science/article/abs/pii/S0048969723043693
  113. https://academic.oup.com/jid/article/221/Supplement_4/S436/5727782
  114. https://conbio.onlinelibrary.wiley.com/doi/10.1111/conl.13069
  115. https://cites.org/sites/default/files/documents/E-AC32-Inf-10.pdf
  116. https://neprimateconservancy.org/lion-tailed-macaque/
  117. https://www.bbc.com/news/science-environment-37490932
  118. https://www.gibraltar.gov.gi/environment/ape-management
  119. https://zslpublications.onlinelibrary.wiley.com/doi/10.1111/acv.70015
  120. https://wanprc.uw.edu/20-years-of-community-outreach-education-in-tangkoko-indonesia/
  121. https://taiwania.ntu.edu.tw/download/tai.2006.51.258.pdf/512/issue
  122. https://www.aph.gov.au/DocumentStore.ashx?id=31c2c61a-5f80-41ca-a43e-55fd305c3f62
  123. https://pmc.ncbi.nlm.nih.gov/articles/PMC9312286/
  124. https://www.itis.gov/servlet/SingleRpt/SingleRpt?search_topic=TSN&search_value=180097
  125. https://bmcgenomics.biomedcentral.com/articles/10.1186/s12864-015-1437-0
  126. https://www.sciencedirect.com/science/article/abs/pii/S1055790322001269
  127. https://pmc.ncbi.nlm.nih.gov/articles/PMC10413639/
  128. https://www.researchgate.net/publication/228727780_Anthropogenic_macaque_hybridization_and_genetic_pollution_of_a_threatened_population
  129. https://www.iucnredlist.org/search?query=Macaca&searchType=species
  130. https://www.sciencedirect.com/science/article/abs/pii/S0047248423001331
  131. https://www.sciencedirect.com/science/article/abs/pii/S0047248417302348
  132. https://ui.adsabs.harvard.edu/abs/2014QSRv...96..117E/abstract
  133. https://www.sciencedirect.com/science/article/abs/pii/S0277379114001565
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