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Old World monkey

Old World monkeys are belonging to the family Cercopithecidae, a diverse group comprising approximately 160 species primarily native to and . They are distinguished from by several key anatomical features, including narrow, downward-facing nostrils, non-prehensile tails, and bilophodont molars adapted for grinding vegetation. The family originated in the during the epoch, around 24–30 million years ago, and represents the most species-rich lineage of catarrhine , with evolutionary adaptations reflecting a range of ecological niches from terrestrial savannas to arboreal forests. Cercopithecidae is divided into two main subfamilies: Cercopithecinae (cercopithecines), which includes omnivorous genera such as macaques (Macaca), baboons (Papio), and guenons (Cercopithecus), and Colobinae (colobines), which consists of folivorous, leaf-eating genera like colobus monkeys (Colobus) and langurs (Presbytis). Cercopithecines typically feature cheek pouches for storing food and exhibit greater terrestrial tendencies, while colobines possess specialized sacculated stomachs for fermenting fibrous plant material, enabling efficient digestion of mature leaves. Physical characteristics common to the family include ischial callosities—hardened skin pads on the buttocks for sitting—and a dental formula of 2.1.2.3, with pronounced sexual size dimorphism in many species, where males can be significantly larger than females. Tail length varies widely, from short in baboons to long in some macaques, but none are prehensile for grasping. These monkeys inhabit a broad spectrum of environments, from tropical rainforests and montane forests to grasslands and semi-deserts, with cercopithecines showing higher diversity in and colobines peaking in Asian forests. Socially, Old World monkeys are highly gregarious, forming complex troops that range from multimale-multifemale groups in macaques to one-male harems in some colobines, with behaviors including grooming, vocalizations, and hierarchical structures that enhance group cohesion and predator defense. Many species, such as the (Macaca mulatta), play significant roles in biomedical due to their physiological similarities to humans, while others face conservation threats from loss and hunting.

Taxonomy and Phylogeny

Classification

Old World monkeys belong to the kingdom Animalia, phylum Chordata, class Mammalia, order Primates, suborder Haplorhini, infraorder Simiiformes, parvorder Catarrhini, superfamily Cercopithecoidea, and family Cercopithecidae. The family Cercopithecidae is the largest within the order Primates, encompassing 24 genera and 160 species distributed across Africa and Asia. This family is divided into two distinct subfamilies: Cercopithecinae and Colobinae, which differ in diet, digestive adaptations, and ecological niches. The Cercopithecinae subfamily includes omnivorous species with cheek pouches that allow temporary food storage, facilitating opportunistic foraging; representative genera include Macaca (macaques, with about 22 species across Asia and North Africa) and Papio (baboons, with five species primarily in Africa). These monkeys are often terrestrial or semi-arboreal and exhibit diverse social structures adapted to varied habitats from savannas to forests. In contrast, the subfamily consists of predominantly folivorous species lacking cheek pouches but possessing complex, sacculated stomachs with chambers for efficient leaf digestion. Key genera include Colobus (colobus monkeys, with five species in African forests) and Trachypithecus (Asian langurs, with around 11 species), which are highly arboreal and rely on specialized microbial symbionts to break down fibrous vegetation. This subfamily emphasizes plant-based diets, with some species selectively browsing on mature leaves or unripe fruits to minimize secondary compounds. Recent studies continue to refine colobine . Taxonomic classifications within Cercopithecidae have undergone significant revisions driven by , particularly in the diverse tribe () of the . Traditionally, many species were grouped under the single genus Cercopithecus, but genomic and museomic analyses have supported splitting them into distinct genera such as Chlorocebus (vervets and grivets), Allenopithecus (Allen’s swamp monkey), and Miopithecus (talapoins), reflecting deeper evolutionary divergences and resolving polyphyletic groupings. These revisions, informed by mitochondrial and nuclear DNA sequences, have contributed to recognizing approximately 40 across six genera, enhancing conservation assessments. Within the parvorder , monkeys (superfamily Cercopithecoidea) are the to Hominoidea (apes and humans), sharing narrow-nostriled features that distinguish them from (Platyrrhini).

Evolutionary History

monkeys, belonging to the superfamily Cercopithecoidea, represent one of the two major lineages within the clade of , alongside the Hominoidea (apes and humans). The divergence between and Platyrrhini () occurred approximately 40–50 million years ago during the late Eocene to early , marking a key event in evolution as ancestral catarrhines dispersed from . Within , the split between Cercopithecoidea and Hominoidea took place around 25–30 million years ago in the , with molecular and fossil evidence indicating an African origin for this basal divergence. The earliest definitive fossils of Old World monkeys come from the genus Victoriapithecus, dated to 15–17 million years ago in the middle of , such as specimens from Maboko Island that exhibit primitive catarrhine cranial features including a short face and bilophodont molars. These fossils provide the oldest evidence of the distinctive dental morphology that characterizes cercopithecoids, bridging earlier primitive catarrhines and more derived forms. Although older potential cercopithecoid-like remains exist from the early (~22 million years ago), Victoriapithecus confirms the establishment of the lineage with key anatomical traits. A pivotal evolutionary in monkeys was the development of bilophodont molars, featuring two transverse lophs per tooth that enhanced shearing and grinding capabilities, particularly for processing tough, fibrous vegetation like leaves and seeds. This dental innovation likely arose in response to dietary shifts during the , enabling exploitation of varied ecological niches in forested and environments. Following the , around 5–10 million years ago, monkeys underwent significant radiation, diversifying into over 130 extant species across and as climates cooled and habitats fragmented. Phylogenetically, Cercopithecoidea forms the sister group to Hominoidea within , with analyses supporting this relationship based on genomic data from nuclear genes. The two main subfamilies, (cheek-pouched monkeys) and (leaf-eating monkeys), diverged approximately 15–20 million years ago in the early to middle , with subsequent radiations reflecting adaptations to folivory and omnivory, respectively. These estimates derive from relaxed clock models calibrated against fossil constraints, highlighting the tempo of cercopithecoid evolution.

Physical Characteristics

Anatomy and Morphology

Old World monkeys, members of the family Cercopithecidae, exhibit a suite of anatomical features adapted to their diverse ecological niches across and . These include a characteristic suited for processing varied material, robust skeletal structures for terrestrial and , and specialized skin modifications for prolonged sitting on hard substrates. The dental formula of monkeys is 2.1.2.3 / 2.1.2.3, consisting of two incisors, one , two premolars, and three molars per , identical to that of apes and humans. This formula supports a ranging from fruits to leaves, with bilophodont molars—featuring two transverse ridges on the occlusal surface—facilitating efficient grinding of fibrous , particularly in folivorous species. Unlike , monkeys possess non-prehensile tails that serve primarily for balance rather than grasping, reflecting adaptations to a mix of quadrupedal and climbing lifestyles. All develop ischial callosities prenatally—thickened, hairless pads of skin over the —that provide cushioning for sitting on rocky or bare ground, a trait unique among anthropoids. Morphological specializations distinguish the two main subfamilies: (cercopithecines) and (colobines). Cercopithecines feature internal cheek pouches for temporary , enabling rapid collection and consumption during , while colobines exhibit thumb reduction or absence, an that enhances leaf-stripping efficiency with their specialized hands. Sexual dimorphism is pronounced in many Old World monkeys, particularly in body size and canine teeth, where males are substantially larger and possess elongated, projecting canines for intra-sexual and display. For instance, in species like baboons, males can weigh twice as much as females, with canines exceeding those of comparably sized carnivores in length. Sensory adaptations include routine trichromatic color vision in all Old World monkeys, achieved through three opsin genes that distinguish , , and wavelengths, an evolutionary innovation shared with apes that aids in detecting ripe fruits and social signals. Compared to apes, Old World monkeys retain enhanced olfactory capabilities, with a higher proportion of functional genes and relatively larger olfactory bulbs, supporting scent-based and communication.

Size and Appearance

Old World monkeys exhibit a wide range of body sizes, from the smallest species, the (Miopithecus talapoin), with a head and body length of 32 to 45 cm and weight of 0.8 to 1.9 kg, to the largest, the (Mandrillus sphinx), where males reach 61 to 76 cm in head and body length and up to 54 kg in weight. This variation spans over an in mass across the family Cercopithecidae, reflecting adaptations to diverse ecological niches. External appearance among Old World monkeys is highly diverse, particularly in fur coloration and facial features. For instance, the (Macaca silenus) features glossy black fur with a prominent golden or silvery mane around the head and shoulders, while colobus monkeys like the (Colobus guereza) display striking black-and-white pelage with long, flowing mantle hair. Facial and rump features further highlight this diversity; baboons (Papio spp.) often have vividly colored naked rumps in , blue, or purple that intensify during displays, and the (Nasalis larvatus) is distinguished by the males' large, pendulous nose that hangs over the upper lip. Fur is typically coarse and not woolly, ranging from olive-gray in mandrills to reddish-brown in some guenons. Sexual dimorphism is pronounced in many species, with males generally larger than females—often by a factor of two in body mass—and exhibiting brighter coloration or more exaggerated features for displays. In the , for example, adult males have intensely multicolored faces and rumps in red, blue, and violet, contrasting with the duller olive-brown pelage of females. Males also possess prominent canines, though this is tied to underlying dental morphology. Tail length varies relative to body size, being longer in arboreal species such as langurs ( spp.), where tails often exceed body length (e.g., 91 cm tail in Hanuman langurs versus 64 cm body), aiding balance in trees, and shorter in more terrestrial forms like baboons, where tails measure about 45 to 71 cm against a body of 50 to 114 cm.

Habitat and Distribution

Geographic Range

Old World monkeys, belonging to the family Cercopithecidae, are native to and , where they occupy a wide array of environments across these continents. In , prominent examples include baboons of the genus Papio, which range widely from savannas to woodlands, and colobus monkeys of the genus Colobus, concentrated in forested regions. In , macaques of the genus Macaca exhibit an extensive distribution from in the north to in the south, while langurs such as those in the genus are prevalent in and . The family comprises approximately 160 (as of 2024), with the majority occurring in and the remainder in ; there are no native populations in the or . Fossil evidence reveals a broader historical range, extending into during the Miocene epoch, as exemplified by Mesopithecus, an early colobine monkey whose remains have been found in sites in dating to around 9-7 million years ago. Today, the only extant Old World monkey population outside Africa and is the (Macaca sylvanus) in , which was introduced, likely by humans during historical trade or military activities. Certain regions highlight endemism within this distribution; for instance, lacks any monkeys, its consisting solely of lemurs that evolved in isolation after the island's separation from . Island endemics are notable in , such as the seven species of s restricted to , including the crested black macaque (Macaca nigra), which is found nowhere else.

Habitat Preferences

Old World monkeys, belonging to the family Cercopithecidae, occupy a wide array of habitats across and , reflecting their ecological versatility. Many species, particularly colobines such as those in the genera Procolobus and Piliocolobus in or Semnopithecus and Trachypithecus in , prefer tropical rainforests where they exploit the dense canopy layers for shelter and foraging. In contrast, cercopithecines like baboons (Papio spp.) thrive in open savannas and grasslands, where they navigate expansive, less vegetated landscapes. Montane forests and highland grasslands also support specialized populations, such as geladas (Theropithecus gelada) in the , which inhabit elevations from approximately 1,800 to 4,200 meters. Adaptations to these habitats vary significantly between subfamilies. Arboreal colobines, including langurs ( spp.), have evolved specialized multi-chambered stomachs and high-cusped molars to efficiently digest fibrous leaves in the humid, forested canopies, allowing them to maintain small home ranges often less than 1 km². Terrestrial species, such as patas monkeys (Erythrocebus patas) in African s, exhibit elongated limbs and a locomotor system enabling speeds up to 55 km/h, facilitating rapid evasion of predators in open grasslands. Baboons demonstrate semi-terrestrial adaptations, combining quadrupedal walking on the ground with climbing abilities to access resources in heterogeneous savanna environments. Microhabitat preferences further refine their ecological niches, with many species selecting areas proximate to water sources like or streams to mitigate dehydration risks, particularly during dry seasons. Altitudinal ranges span from in coastal forests to high elevations exceeding 4,000 meters for species like the (Rhinopithecus roxellana) in Asian montane forests. Climate influences habitat choice, with a general preference for warm, humid conditions in tropical zones, though some, like geladas and certain macaques, tolerate cooler, seasonal climates in highlands by adjusting behaviors such as cliff-sleeping for protection or shifting to lichen-based diets in winter.

Behavior

Social Systems

Old World monkeys, belonging to the family Cercopithecidae, typically live in cohesive social groups characterized by matrilineal organization, where adult females and their offspring form the stable core of the troop, while males disperse from their natal groups at to avoid and competition. This female results in kin-based bonds that structure much of the group's dynamics, with females remaining in their birth groups throughout life. Group sizes vary widely depending on species and habitat; for instance, colobine monkeys like the often form troops of 25-50 individuals, whereas hamadryas baboons (Papio hamadryas) aggregate into larger multilevel societies exceeding 200 members, consisting of one-male units clustered into clans and bands. Dominance hierarchies within these groups are typically linear and based on , , , and individual attributes such as and , influencing access to resources and opportunities. In multi-male groups, such as those of baboons (Papio cynocephalus), males form coalitions to challenge higher-ranking individuals, enhancing their status and through cooperative alliances that target rivals during conflicts. Female hierarchies are similarly matrilineal, with inherited from mothers, promoting nepotistic support among kin in grooming and agonistic interactions. Males employ various strategies post-dispersal, including forming all-male groups on the of troops or living solitarily until they can for entry into a breeding unit. In species like savanna baboons, incoming males may commit against unrelated infants to shorten female interbirth intervals and accelerate their return to estrus, thereby accelerating the males' own reproductive opportunities. This tactic is observed across multiple cercopithecid species, including langurs and macaques, where it aligns with male takeover events in unstable groups. In larger troops, —non-maternal care provided by other females—plays a key role in survival, with unrelated or females carrying, grooming, and protecting young to alleviate maternal burden and enhance group cohesion. For example, in golden snub-nosed monkeys (Rhinopithecus roxellana), over 87% of receive allonursing from additional adult females, which supports faster and reduces predation risk in complex social settings. This cooperative care is particularly adaptive in matrilineal societies, fostering reciprocal benefits among females.

Locomotion and Communication

Old World monkeys display a range of styles adapted to their terrestrial or arboreal lifestyles. Terrestrial , such as baboons, primarily employ quadrupedal walking and galloping on the ground and along branches, often adopting hand postures to enhance stride length and stability during rapid movements. Arboreal forms, like langurs, favor leaping between supports and climbing on slender substrates, with leaps predominating on twigs and lianas to navigate fragmented forest canopies efficiently. Colobine monkeys, including odd-nosed , frequently use suspensory and vertical climbing, suspending from all four limbs to access foliage in dense vegetation, reflecting adaptations for their folivorous diet and larger body sizes. Most monkeys are diurnal, active primarily during daylight hours and retiring to sleeping sites at night to minimize predation risk. They select elevated sites such as tree crowns, cliffs, or rock faces for , often reusing preferred locations for protection from ground-dwelling predators and inclement weather. use remains rare among monkeys, though long-tailed macaques (Macaca fascicularis) in coastal habitats employ stone hammers and anvils to crack shellfish and nuts, producing archaeological signatures similar to those of early hominins but limited to specific populations. Communication in Old World monkeys encompasses vocal, visual, and olfactory modalities, enabling coordination and predator avoidance. Vocalizations include predator-specific alarm calls, as in vervet monkeys (Chlorocebus pygerythrus), where distinct acoustic signals for aerial, terrestrial, or snake threats elicit tailored escape behaviors like looking up, seeking cover, or standing bipedally. Facial expressions convey affiliation and intent; for instance, lip-smacking in macaques involves rhythmic jaw oscillations that signal social tolerance and may parallel proto-speech rhythms in . Olfactory signals involve marking and genital rubbing to deposit scents, with males and females using these to advertise dominance, reproductive , or boundaries, particularly in species like Japanese macaques (Macaca fuscata). These diverse signals collectively support social cohesion by facilitating interactions and group vigilance.

Ecology

Diet and Foraging

Old World monkeys display varied dietary compositions shaped by their ecological niches, with the two main subfamilies exhibiting distinct feeding habits. Colobines () are predominantly folivorous, relying heavily on leaves, young shoots, and seeds, often comprising over 90% of their intake in species like the red colobus (Piliocolobus tephrosceles), where leaves account for approximately 92% of the diet. In contrast, cercopithecines (), such as baboons (Papio spp.) and macaques (Macaca spp.), are omnivorous and opportunistic, consuming a broad range including fruits, seeds, flowers, buds, , and occasionally small vertebrates, with baboons documented eating up to 69 food items from 29 species over a 30-day period. These differences reflect adaptations to resource availability, with colobines favoring fibrous, low-quality foliage in forested habitats and cercopithecines exploiting diverse, patchier resources in more open environments. Foraging techniques vary by habitat and diet, often involving group-based strategies to access food sources efficiently. Arboreal colobines, such as the (Nasalis larvatus), employ canopy stripping with specialized hands to harvest leaves and unripe fruits from tree branches, while spending significant time in the upper forest layers. Terrestrial cercopithecines like s forage on the ground in troops, digging for , tubers, and underground storage organs using their robust hands and opportunistic predation on during dry seasons when plant matter is scarcer. Seasonal shifts are common across both subfamilies; for instance, colobines like the maroon leaf monkey (Presbytis rubicunda) increase seed consumption during fruiting periods, while snub-nosed monkeys (Rhinopithecus spp.) turn to lichens in winter when foliage is limited. Digestive adaptations enable efficient processing of these diets, with colobines featuring a multi-chambered for rumination-like of fibrous leaves by symbiotic microbes, as seen in the proboscis monkey's sacculated stomach that detoxifies plant defenses and extracts nutrients from high-fiber foods. Cercopithecines, adapted for rapid , possess pouches to store food for quick consumption during foraging bouts, allowing them to handle a mix of easily digestible fruits and tougher items like seeds and . Resource competition influences patterns, particularly through territorial behaviors defending key food sites. Male guerezas (Colobus guereza) mediate intergroup encounters to protect fruiting trees and fallback resources like during , reducing within groups. Fallback foods, such as lichens for snub-nosed monkeys or underground tubers for s, sustain populations when preferred items decline seasonally, highlighting the role of dietary flexibility in survival.

Reproduction and Life Cycle

Old World monkeys, belonging to the Cercopithecidae, typically exhibit polygynandrous systems within multi-male, multi-female groups, where females with multiple males during their estrous periods to reduce risk and enhance . In many species, such as baboons (Papio spp.) and macaques (Macaca spp.), male-male competition for access to receptive females influences success, while female choice plays a significant role in partner selection. Breeding is often seasonal in temperate or high-altitude populations, synchronizing with resource availability; for instance, macaques (Macaca fuscata) primarily from October to February during winter months. Gestation periods in Old World monkeys generally last 5 to 7 months, varying by —for example, approximately 164 days in rhesus macaques (Macaca mulatta) and 173 days in Japanese macaques. Births typically produce a single , with twinning being rare at rates of about 0.18% to 0.21% across like macaques. Newborns weigh roughly 10% to 20% of the mother's body mass, such as 400–500 grams for infants of female rhesus macaques weighing 4–8 kilograms, enabling initial ventral carriage by the mother. Infants exhibit high dependency on their mothers for the first 6 to 12 months, during which they are carried, nursed, and protected, with carriage shifting from ventral to dorsal after 2–3 months in species like baboons. Weaning occurs gradually between 1 and 2 years, allowing juveniles to remain socially affiliated with their mothers post-weaning. Allomaternal care, including nursing and grooming by non-mothers, supplements maternal efforts in some species, such as golden snub-nosed monkeys (Rhinopithecus roxellana), where it occurs routinely in the early months to support infant survival. Sexual maturity is reached at 3–5 years for females, often earlier than males at 4–8 years, marking the transition to reproductive adulthood. In the wild, monkeys have lifespans of 10–30 years, influenced by predation, , and resource scarcity, though individuals in can live up to 40 years due to veterinary care and stable . Reproductive output peaks in mid-adulthood, with females capable of bearing offspring from ages 4 to 18, after which declines. This underscores the ' investment in prolonged to ensure offspring survival in complex social environments.

Conservation and Human Relations

Threats and Conservation Status

Old World monkeys face significant threats from habitat loss, primarily driven by deforestation for agriculture, logging, and urbanization across their native ranges in and Asia. This anthropogenic pressure affects a substantial portion of the family Cercopithecidae, with identified as the leading cause of decline for many . Hunting poses another major threat, including for consumption and , particularly impacting species like macaques in and various colobines in . For instance, long-tailed macaques (Macaca fascicularis) have experienced severe pressure for these purposes, contributing to rapid population reductions. According to the , approximately 72% of the 138 recognized Old World monkey species are classified as threatened with extinction (Vulnerable, Endangered, or ) as of 2024, with 26 species in the category. Miss Waldron's red colobus (Piliocolobus waldroni), for example, is listed as and possibly extinct due to combined loss and hunting in its restricted range in and Côte d'Ivoire. A notable number of species remain , often because their remote forest habitats limit comprehensive surveys and population assessments. As of the 2023–2025 " in Peril" report, nearly two-thirds of species, including many Old World monkeys, are threatened, with ongoing assessments like the reaffirmation of Endangered status for long-tailed macaques in 2024. Emerging threats include , which is projected to alter suitability and structure in tropical forests, exacerbating range contractions for many monkeys. Disease transmission, such as virus outbreaks in species, further endangers populations by causing high mortality in susceptible groups like colobines and cercopithecines. Population trends indicate sharp declines across several genera, with some experiencing reductions of up to 50% over the past 30 years due to intensified hunting and . In introduced areas, such as parts of where rhesus macaques (Macaca mulatta) have established invasive populations, competition with native wildlife adds pressure to local ecosystems.

Interactions with Humans

Old World monkeys hold significant cultural roles in various societies, particularly in where gray langurs (Semnopithecus entellus), also known as Hanuman langurs, are revered as sacred due to their association with the monkey god from the epic. This reverence often leads to protection and feeding by devotees, influencing local human-monkey interactions in . In biomedical research, rhesus macaques (Macaca mulatta) have been extensively used as models for studying human diseases, contributing to advances in , infectious diseases, aging, and reproductive health. Studies on (SIV) in sooty mangabeys (Cercocebus atys), natural hosts that do not progress to AIDS-like illness, have provided key insights into HIV pathogenesis and development by revealing mechanisms of non-pathogenic infection. However, ethical concerns surrounding the use of monkeys in research include welfare issues related to , invasive procedures, and the moral implications of utilizing phylogenetically close species, prompting exploration of alternatives such as genetically modified or computational models. Human-monkey conflicts arise from resource competition, notably crop raiding by chacma baboons (Papio ursinus) in South African farmlands adjacent to protected areas, where they are reported as the primary raiders, leading to economic losses for farmers. In urban settings like , , rhesus macaques have adapted to environments, exploiting food waste and provisioning, which exacerbates conflicts through and health risks from close proximity to humans. Conservation initiatives for Old World monkeys include protected areas in the , such as the Wildlife Reserve, which safeguards species like mangabeys and mandrills amid broader efforts. Reintroduction programs, exemplified by the Golden Langur Conservation Project in , have successfully increased populations of Gee's golden langurs (Trachypithecus geei) from around 1,500 in 1997 to over 6,000 by 2024 through habitat restoration and translocation. Additionally, regulates international trade in vulnerable Old World monkeys, listing species such as Diana monkeys (Cercopithecus diana), lion-tailed macaques (Macaca silenus), and (Mandrillus leucophaeus) in Appendix I to prohibit commercial trade, while others like mantled guerezas (Colobus guereza) fall under Appendix II for monitored exports.

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