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Colobinae

Colobinae is a subfamily of Old World monkeys in the family Cercopithecidae, consisting of approximately 10 genera and over 60 species of arboreal primates specialized in folivory, commonly known as leaf monkeys, colobus monkeys, and their relatives.^[1] These monkeys are distinguished by their complex, multi-chambered stomachs that house symbiotic microbes for fermenting cellulose-rich plant material, high-cusped molars for grinding tough foliage, and the absence of cheek pouches found in other cercopithecids.^[1]^[2] Native to Africa and Asia, colobines inhabit a wide range of forested environments, from tropical lowlands and mangroves to montane forests up to 4,700 meters in elevation, including peat swamps, limestone karsts, and riparian zones.^[1] Their distribution spans sub-Saharan Africa—from Senegal to Ethiopia and Zanzibar—and southern Asia, from Sri Lanka and India to southern China, Indonesia.^[1] African genera include Colobus (five species of black-and-white colobuses), Piliocolobus (17 species of red colobuses)^[3], and Procolobus (one species, the olive colobus), while Asian genera encompass Semnopithecus (gray langurs), Trachypithecus (leaf monkeys), Presbytis (surilis), Pygathrix (douc langurs), Rhinopithecus (snub-nosed monkeys), Nasalis (proboscis monkey), and Simias (pig-tailed langur).^[1]^[4] Colobines are primarily folivorous, with leaves comprising up to 92% of the diet in some species like red colobuses, supplemented seasonally by fruits, seeds, flowers, and occasionally lichens or bark.^[1] Their slender builds and long tails facilitate arboreal locomotion, and social structures vary from unimale-multifemale groups in many species to multimale-multifemale troops in others, often featuring territorial behaviors and male dispersal at maturity.^[2] Many colobine species face conservation threats, with several classified as endangered or vulnerable due to habitat loss and hunting.^[1]

Taxonomy and Phylogeny

Classification

Colobinae constitutes a subfamily within the family Cercopithecidae, encompassing the folivorous Old World monkeys that diverged from the sister subfamily Cercopithecinae approximately 15–20 million years ago.^[5] The subfamily is currently classified into two primary tribes: Colobini, which includes the African colobines represented by the genera Colobus, Piliocolobus, and Procolobus; and Presbytini, comprising the Asian langurs such as Trachypithecus, Semnopithecus, and Rhinopithecus.^[6]^[7] Colobinae encompasses approximately 78 species distributed across 10 genera, with notable examples including the mantled guereza (Colobus guereza) among the African taxa and the proboscis monkey (Nasalis larvatus) within the Asian group.^[8] Taxonomic revisions in recent decades, driven by molecular phylogenetic analyses, have refined this structure, including the recognition of the odd-nosed monkeys—encompassing genera Nasalis, Simias, Pygathrix, and Rhinopithecus—as a monophyletic clade within the Presbytini tribe.^[9]

Evolutionary History

The subfamily Colobinae, comprising leaf-eating monkeys, has a fossil record extending back to the middle Miocene epoch, approximately 15 million years ago, with initial discoveries in Africa and subsequent dispersals into Asia. Primitive colobine fossils from middle to late Miocene sites in Africa, such as those from the Ngorora Formation in Kenya, indicate an African origin for the group.^[5]^[10] In Asia, significant late Miocene deposits from the Siwalik Hills in northern India and Pakistan yield colobine remains dating to around 9–7 million years ago, including species like Mesopithecus sivalensis, which suggest early diversification and adaptation to forested environments across Eurasia.^[11] These fossils document a gradual expansion from African cradle sites to Asian continental interiors, facilitated by tectonic uplifts and climatic shifts during the Miocene.^[12] Phylogenetic reconstructions based on both morphological and molecular data affirm Colobinae as a monophyletic clade within the Cercopithecidae family, with a foundational divergence separating the African (e.g., Colobina) and Asian (e.g., Presbytina) lineages estimated at 12-15 million years ago.^[13] This basal split is supported by analyses of complete mitochondrial genomes and nuclear markers, which place the common ancestor of modern colobines in the mid-Miocene, followed by vicariant events tied to the closure of the Tethys Sea and the emergence of the Himalayan barrier.^[14] Within these lineages, further cladogenesis occurred, with Asian colobines exhibiting monophyly in the odd-nosed clade, underscoring the complexity of post-Miocene radiations.^[9] Molecular genetic studies, particularly those employing mitochondrial DNA sequences, reveal a phase of rapid radiation among colobine taxa during the Pliocene epoch (5.3-2.6 million years ago), coinciding with global cooling, aridification, and episodic forest fragmentation across Africa and Asia.^[15] These environmental pressures likely promoted speciation through isolation in refugia, as evidenced by divergence estimates for genera like Trachypithecus and Pygathrix around 4-3 million years ago.^[16] Such dynamics are corroborated by phylogeographic models integrating fossil calibrations, highlighting how habitat mosaics drove lineage sorting and endemism.^[17] This evolutionary trajectory reflects an adaptive radiation centered on folivory, where colobines diverged from more frugivorous cercopithecoid ancestors by developing specialized dentition, including high-crowned molars and bilophodont cusps optimized for processing fibrous leaves and unripe fruits.^[18] Post-divergence refinements in cranio-dental morphology, observed in Miocene fossils transitioning to Pliocene forms, enabled exploitation of low-nutrient, high-fiber diets in fragmented woodlands, distinguishing colobines from sympatric frugivores.^[19] This dietary specialization, evolving concurrently with rumen-like forestomach fermentation, underpinned their ecological success and diversification.^[20]

Hybrids

Hybridization within the Colobinae subfamily is rare but documented in both natural and captive settings, often resulting from habitat overlap or human-mediated encounters, and it raises questions about species boundaries, particularly in the tribe Presbytini. In African colobines, natural hybrid zones occur where ranges overlap, such as between the king colobus (Colobus polykomos) and ursine colobus (Colobus vellerosus) in West Africa, where the taxon Colobus polykomos dollmani represents a hybrid swarm with predominant C. vellerosus genetic contributions due to secondary contact in fragmented forests between the Bandama and Sassandra rivers.^[21] These hybrids exhibit intermediate morphological traits, illustrating how habitat fragmentation can facilitate gene flow between closely related species. Although the outline suggested hybrids between Colobus guereza and C. polykomos, their non-overlapping ranges (eastern vs. western Africa) preclude natural occurrences, but captive breeding has produced such hybrids in zoos, as recorded in early studies.^[22] In Asian colobines, natural hybridization has been increasingly documented amid habitat loss, particularly within the genus Trachypithecus of the Presbytini tribe. For instance, mixed-species groups of Phayre's langur (Trachypithecus phayrei) and capped langur (T. pileatus) in northeast Bangladesh have led to genetically confirmed hybrid offspring, driven by forest fragmentation forcing sympatry in areas like Satchari National Park. Captive hybridization between Trachypithecus species, such as T. phayrei and T. pileatus, has also occurred in zoos, though less frequently reported; these often face fertility challenges stemming from chromosomal rearrangements, with Trachypithecus species showing variations in karyotypes (e.g., 44 chromosomes but differing arm numbers) that can cause meiotic issues in hybrids.^[8] Another example involves a putative intergeneric hybrid between the proboscis monkey (Nasalis larvatus) and silvery langur (Trachypithecus cristatus) in Borneo's Lower Kinabatangan Wildlife Sanctuary, resulting from habitat overlap in degraded mangroves.^[23] Genetic markers, including microsatellite loci and mitochondrial DNA analysis, have been crucial for confirming hybrid ancestry in colobines by detecting mixed parental contributions. For example, microsatellite genotyping of fecal DNA from T. phayrei × T. pileatus hybrids revealed heterozygous profiles with alleles from both species, supporting ongoing gene flow.^[24] Similarly, nuclear and mitochondrial discrepancies in colobine populations have evidenced ancient hybridization events, complicating phylogenetic reconstructions. These findings imply taxonomic revisions in Presbytini, where hybrid zones suggest porous species boundaries and potential merging of lineages under anthropogenic pressures, challenging traditional classifications based on morphology alone.^[25]

Morphology and Adaptations

Physical Characteristics

Colobinae species exhibit a wide range of body sizes, typically spanning 4 to 24 kg across the subfamily, with adults in smaller genera like Colobus weighing around 5-10 kg and larger Asian forms such as the proboscis monkey (Nasalis larvatus) reaching up to 20 kg in males.^[7] Sexual dimorphism is pronounced in body mass for most species, with males generally 20-100% heavier than females; for instance, proboscis monkey males average 19.8 kg compared to 9.6 kg for females, reflecting extreme dimorphism linked to social and ecological factors.^[7]^[26] The pelage of Colobinae is dense and varies markedly by genus and region, often featuring combinations of black, white, gray, olive, or reddish hues, with some Asian species displaying more vibrant or silvery tones in juveniles that fade with age.^[7] Ischial callosities, the hardened skin pads on the buttocks typical of Old World monkeys, are prominent and colorful (often pink or red) in many Colobinae, aiding in prolonged sitting on branches.^[27] Certain genera, such as Nasalis and Pygathrix, possess expandable throat sacs or laryngeal sacs that amplify vocalizations, particularly in males during displays.^[28] Limb proportions in Colobinae are adapted for an arboreal lifestyle, with elongated hindlimbs relative to forelimbs (intermembral indices often below 80) supporting quadrupedal locomotion, leaping, and suspensory behaviors like brachiation in forested environments.^[28] Forelimbs are slender and flexible, facilitating grasping and suspension, while tails are typically long and non-prehensile, providing balance during movement.^[7] Thumbs are reduced or absent in many species, enhancing hook-like grips for foliage navigation.^[7] Cranial morphology in Colobinae features a relatively short but prognathic muzzle with narrow incisors and a broad interorbital region, adaptations that accommodate their folivorous diet through specialized dentition briefly referenced here as high-cusped molars for leaf processing.^[7] Eye orbits are large and forward-facing, enhancing binocular vision in dim understory conditions, though most species are diurnal rather than strictly nocturnal.^[7]^[9]

Specialized Traits

Colobinae exhibit a highly specialized digestive system adapted for folivory, featuring a complex, multichambered forestomach that facilitates microbial fermentation of fibrous plant material. The forestomach is divided into the saccus gastricus (with or without a distinct presaccus) and the tubus gastricus/pars pylorica, where symbiotic bacteria break down cellulose and hemicellulose into volatile fatty acids under near-neutral pH conditions (5.0–7.0), enabling efficient nutrient extraction from leaves that would otherwise be indigestible.^[29] In the tribe Colobini, the stomach is typically sacculated into a tripartite structure (lacking a clear presaccus) or quadripartite form (including the presaccus), which enhances fermentation capacity and supports high-fiber diets comprising up to 52% acid detergent fiber.^[29] The dentition of Colobinae is finely tuned for processing tough foliage, with a permanent dental formula of 2.1.2.3 (two incisors, one canine, two premolars, and three molars per quadrant), shared with other Old World monkeys but specialized in the molars.^[30] The molars are high-crowned and exhibit bilophodonty, characterized by four cusps connected by prominent anterior and posterior transverse crests that form shearing surfaces ideal for grinding and fragmenting fibrous leaves.^[30] Lower third molars often include well-developed hypoconulids, further aiding in the mastication of abrasive plant matter, while the overall dental arcade supports heavy chewing pressures without the need for extensive prognathism.^[30] Sensory adaptations in Colobinae reflect a shift away from olfactory reliance, with the highest rates of olfactory receptor gene loss observed among primates, particularly in this subfamily, potentially linked to their folivorous lifestyle and reliance on visual cues.^[31] Compared to Cercopithecinae, Colobinae show further reduction in olfactory bulb size relative to endocranial volume, a trend that evolved post-15 million years ago in cercopithecoids but accelerated in leaf-eating lineages, compensating through enhanced visual and vocal communication for social and foraging needs.^[32] Additional anatomical specializations include enlarged salivary glands, notably in langurs (tribe Presbytini), which produce tannin-binding proteins to neutralize plant secondary compounds like tannins that inhibit protein digestion and act as toxins in leafy diets.^[29] These proteins, often proline-rich, precipitate tannins in the mouth, preventing their interference with foregut fermentation and allowing access to nutrient-rich foliage.^[29] In some species, such as those in the genus Colobus, the thumb is reduced to a small tubercle or absent, streamlining hand morphology for efficient quadrupedal progression and leaping in arboreal environments.^[33] This reduction, evident in fossil records as early as 1.9 million years ago, distinguishes African colobines from their Asian counterparts and enhances locomotor stability during pronograde movement.^[33]

Distribution and Ecology

Geographic Range

The subfamily Colobinae, comprising leaf monkeys and colobus monkeys, exhibits a broad but disjunct distribution across sub-Saharan Africa and Asia. In Africa, colobines are primarily found in tropical regions from Senegal in the west to Ethiopia in the east, extending southward to Angola and Tanzania. The genus Colobus, including black-and-white and black colobus species, occupies a wide range from Guinea-Bissau through Central Africa to Tanzania and Ethiopia, often south of the Congo River. Red colobus monkeys (Piliocolobus spp.) show similar breadth, with distributions spanning West Africa from Senegal to Ghana, Central Africa including the Congo Basin, and East Africa up to Kenya, Tanzania, and Zanzibar. The Congo Basin stands out as a hotspot of diversity, hosting multiple sympatric Piliocolobus species such as P. bouvieri, P. tholloni, and P. oustaleti, alongside hybrids and co-occurring Colobus taxa like C. satanas and C. guereza.^[34]^[35] In Asia, Colobinae occupy a continuous but fragmented range from Sri Lanka and India through Southeast Asia to southern China, encompassing insular populations on Borneo, Java, Sumatra, and smaller Sundaic islands. The langur genus Semnopithecus is prevalent in South Asia, with subspecies like S. entellus extending from central India to northwestern Bhutan and possibly Afghanistan. Trachypithecus species, numbering around 20, dominate Southeast Asian distributions, occurring in Bhutan, Assam (India), Bangladesh, Myanmar, Thailand, Cambodia, Laos, Vietnam, and southern China, while also inhabiting the Malay Peninsula and Indonesian islands including Borneo and Java. Other genera, such as Presbytis and Rhinopithecus, further diversify the Asian range, with odd-nosed colobines like the proboscis monkey (Nasalis larvatus) restricted to Borneo.^[36]^[16] This African-Asian disjunction reflects ancient biogeographic patterns, with Colobinae originating in Africa during the Early Miocene (approximately 18-16 million years ago) and dispersing to Eurasia via an emerging land bridge between Africa and the Arabian Peninsula. Subsequent tectonic and climatic changes isolated the two continental populations, resulting in independent radiations without subsequent colonization of Madagascar (home to strepsirrhine primates) or Australia. No Colobinae occur in these regions due to historical barriers like the Mozambique Channel and Wallace's Line.^[15] Contemporary distributions have undergone significant contractions, driven primarily by deforestation, with forest cover in primate ranges declining sharply over the 20th century. Across Colobinae habitats, habitat loss has reduced suitable areas by substantial margins, exemplified by the golden langur (Trachypithecus geei) in Assam, India, where only about 18.5% of its potential 66,320 km² range remains suitable due to fragmentation and agricultural expansion. Overall, 96% of African colobine species face extinction risks tied to such losses, underscoring the vulnerability of their ranges since the early 1900s.^[37]^[35]^[38]

Habitat Preferences

Colobinae species predominantly occupy tropical and subtropical forest ecosystems across sub-Saharan Africa and South and Southeast Asia, favoring environments that support their folivorous diets through abundant foliage. These habitats range from lowland primary rainforests to montane forests, with many taxa exhibiting a strong preference for closed-canopy systems that provide structural complexity for locomotion and resource access. For instance, African colobines such as black-and-white colobus (Colobus guereza) thrive in moist evergreen and semi-deciduous forests, while Asian presbytines like purple-faced langurs (Semnopithecus vetulus) utilize a broader spectrum including dry deciduous and coastal forests.^[35]^[7] Altitudinal distribution varies widely among colobines, spanning from sea level to elevations exceeding 4,000 meters, allowing adaptation to diverse climatic gradients. Species like the golden snub-nosed monkey (Rhinopithecus roxellana) inhabit coniferous and mixed broadleaf-conifer forests in the Himalayan foothills at altitudes of 1,500–4,000 meters, where cooler temperatures and seasonal snowfall influence their ranging patterns.^[39] Most colobines lead strictly arboreal lives, selecting microhabitats within the upper canopy layers—typically 20–30 meters above ground—where mature leaves and young shoots are plentiful, directly influencing group ranging distances based on leaf phenology and patchiness. Folivore-specialized taxa, such as red colobuses (Piliocolobus spp.), concentrate activities in the main and emergent canopy strata of primary rainforests, minimizing time in understory or ground levels to optimize energy efficiency. Seasonal variations in resource availability prompt adjustments in movement; for example, some Asian langurs, including Thomas's langur (Presbytis thomasi), exhibit expanded daily ranges or localized shifts during fruiting peaks in subtropical dry forests, though full migrations are rare compared to frugivores.^[40]^[7]^[41]

Behavior and Sociality

Colobine monkeys, a subfamily of Old World monkeys, form a variety of social groups, including multimale-multifemale troops, unimale-multifemale units, and multilevel societies that range in size from 10 to over 300 individuals, with females typically outnumbering males to support cohesive unit stability.^[7] These groups facilitate collective defense against predators and resource sharing, though structures vary across species; for instance, African colobines like the Angola colobus (Colobus angolensis) organize into multilevel societies comprising one-male reproductive units and associated multimale bachelor bands, allowing for fluid male dispersal and reduced infanticide risk.^[42] In Asian colobines such as Hanuman langurs (Semnopithecus entellus), groups often include multiple adult males who compete for mating access, contributing to dynamic social bonds within the troop.^[43] Dominance hierarchies within colobine groups are generally linear and based on factors like age, body size, and tenure, with older or larger individuals holding higher ranks to regulate access to food and mates.^[44] These hierarchies are enforced through agonistic displays rather than frequent physical aggression, including dramatic leaps through the canopy and vocal threats that minimize energy expenditure while asserting status.^[45] In species like the black-and-white colobus (Colobus guereza), age-graded male hierarchies emerge, where subordinates defer to seniors via submissive postures, promoting group harmony amid limited resources.^[46] Communication in colobine societies relies heavily on vocalizations and olfactory signals to maintain cohesion and boundaries. Loud roars and whoops, as seen in gray langurs (Semnopithecus spp.), function primarily for territorial defense during intergroup encounters, signaling group presence over long distances without direct confrontation. Olfactory marking via urine deposition reinforces these boundaries and individual identity, particularly among males patrolling group peripheries.^[47] Such multimodal signaling helps coordinate group movements and alert members to threats. Allomothering behaviors are widespread among colobine females, where non-mothers actively carry, groom, and protect infants, thereby reducing maternal burden and strengthening female alliances for overall group cohesion.^[48] This cooperative care, observed in species like the capped langur (Trachypithecus pileatus), enhances infant survival rates and fosters reciprocal bonds, as participating females may gain future support in conflicts.^[49] Mating primarily occurs within these stable groups, aligning with the social structure's emphasis on internal affiliations.^[50]

Diet and Foraging

Colobinae, a subfamily of Old World monkeys, exhibit a predominantly folivorous diet, with leaves comprising 50-90% of their annual intake across species.^[51]^[52] This foliage-heavy regimen is supplemented by fruits, seeds, flowers, and occasionally insects, allowing nutritional flexibility in varying habitats.^[53] Their diet often features low protein levels relative to fiber, which is managed through specialized foregut fermentation enabling slow digestion and efficient nutrient extraction from fibrous material.^[51]^[7] Foraging in Colobinae involves selective feeding strategies to maximize nutritional value while minimizing risks from plant defenses. Individuals preferentially consume young leaves, which offer higher protein-to-fiber ratios and greater digestibility compared to mature foliage, thereby avoiding high levels of toxins and digestion inhibitors found in less preferred plant parts.^[54]^[53] Groups typically travel daily path lengths of 1-2 km, covering home ranges methodically to access patches of high-quality forage without excessive energy expenditure.^[55]^[56] Seasonal variations in resource availability prompt dietary shifts among Colobinae, particularly in species like Presbytis, where frugivory increases during dry seasons to exploit unripe fruits and other available supplements when young leaves decline.^[41]^[57] This adaptability helps maintain energy balance amid fluctuating food quality and abundance. Anti-predator behaviors during foraging in Colobinae include vigilance postures, such as upright scanning while feeding, which enhances detection of threats but reduces feeding intake rates as individuals alternate between consumption and monitoring.^[58]^[59] These strategies underscore the trade-offs between resource acquisition and survival in predator-rich environments.

Reproduction and Development

Mating Systems

Colobine monkeys predominantly exhibit polygynous mating systems, characterized by one-male, multi-female social units where the resident male monopolizes access to breeding females within his group. This structure is widespread across the subfamily, including in species like the proboscis monkey (Nasalis larvatus), where dominant males actively defend harems of females against intruders from bachelor groups or rival males, ensuring exclusive mating opportunities.^[60]^[61] In many colobine species, such as langurs and colobus monkeys, these one-male units (OMUs) form the core of larger modular societies, with males aggressively repelling challengers to maintain reproductive control.^[62] Mate selection in colobines involves female preferences for dominant males, often signaled through vocalizations, displays, or physical traits that indicate status and protective ability. For instance, in snub-nosed monkeys (Rhinopithecus spp.), females show attraction to males with sexually selected lip coloration that advertises group-holding dominance, facilitating mate choice within polygynous units.^[63] Incoming males frequently employ infanticide as a strategy to eliminate dependent offspring sired by previous residents, thereby shortening female interbirth intervals and hastening their return to estrus; this tactic is documented in species like red colobus (Piliocolobus spp.) and black-and-white colobus (Colobus guereza), where takeovers by new males lead to targeted killings of infants under six months old.^[64]^[28] Females counter this risk by associating closely with protective resident males or dispersing to safer groups.^[65] Copulatory behavior in colobines typically features brief mounts lasting seconds to minutes, often performed in arboreal settings to minimize predation risk, with most species exhibiting concealed ovulation that obscures the timing of peak fertility. This leads to extended female receptivity, where copulations occur promiscuously throughout the cycle and sometimes during pregnancy, as observed in black-and-white colobus where females solicit over 50% of matings in the days surrounding ovulation.^[66] Interbirth intervals generally span 2-3 years, varying with ecological factors such as resource availability; for example, in red colobus, abundant fruit resources correlate with shorter intervals and higher infant survival, while scarcity prolongs them.^[67]^[68] Infanticide events can further compress these intervals by 6-12 months in affected groups.^[69]

Life Cycle Stages

Colobinae exhibit a gestation period typically ranging from 5 to 7 months, with single births being the norm across the subfamily; twins are rare, particularly in the Colobini tribe, where they have been documented only occasionally in wild populations.^[70] For example, in mantled guerezas (Colobus guereza), gestation lasts approximately 170 days, resulting in one offspring after an interbirth interval of about 22 months.^[71] This reproductive strategy aligns with the subfamily's folivorous diet and arboreal lifestyle, prioritizing fewer, larger offspring to maximize survival in resource-limited environments.^[72] Newborn colobines are born precocial, often fully furred, and enter an initial clinging phase where they maintain close ventroventral contact with the mother for the first 3 to 6 months, relying on her for transport and nursing.^[73] Maternal care is intensive during this period, with infants spending over 90% of their time in contact during the first weeks, gradually decreasing as independence develops; allomaternal care, such as brief carrying by other females, occurs in some species like black-and-white colobuses, aiding social integration.^[74] Weaning typically occurs around 12 to 21 months, varying by species and habitat—for instance, in white-headed langurs (Trachypithecus leucocephalus), it extends to 19–21 months in the wild due to nutritional demands of a leafy diet.^[73] Following weaning, juveniles engage in play behaviors, peaking between 1 and 3 years, which facilitate motor skills, social learning, and foraging proficiency essential for arboreal navigation.^[73] Sexual maturity is reached at 3–5 years for females and 4–7 years for males, with age at first birth often around 4 years in species like gray langurs (Semnopithecus entellus).^[70] In the wild, lifespan averages 15–25 years, though it can exceed 30 years in captivity; for example, Angolan colobuses (Colobus angolensis) live up to 20 years in natural habitats.^[75] Juvenile mortality is high, often 30–50% in the first year, primarily due to predation by raptors, leopards, or chimpanzees, and accidents such as falls from trees during early independence attempts.^[76] In Tana River red colobuses (Piliocolobus rufomitratus), infant mortality reaches 55% within the first year, underscoring the vulnerabilities of this developmental stage.^[76]

Conservation

Major Threats

Habitat destruction, primarily through deforestation for agriculture, logging, and human settlement, poses the most significant threat to Colobinae populations across their ranges in Africa and Asia. In Africa alone, forest loss affects 96% of the 24 colobine species, rendering them threatened with extinction according to IUCN assessments, as these folivorous primates depend on mature forest canopies for their specialized diets.^[35] For instance, the red-shanked douc langur (Pygathrix nemaeus) in Vietnam has experienced severe population declines due to extensive logging and agricultural expansion, contributing to its Critically Endangered status.^[77] Hunting for bushmeat and the pet trade further exacerbates declines, particularly in regions with high human population density and limited enforcement of wildlife laws. This activity is the second major threat to African colobines, impacting 68% of subspecies and driving six species to Critically Endangered levels.^[35] In Central Africa, colobine species like the Central African red colobus (Piliocolobus oustaleti) are heavily targeted, with the bushmeat trade resulting in thousands of primate deaths annually across markets in the region, including colobines.^[78] Climate change disrupts Colobinae habitats by altering rainfall patterns and leaf phenology, which directly affects the availability and nutritional quality of their primary food source—young leaves. Projections indicate that warming and reduced precipitation could decrease leaf protein content and force dietary shifts, with modeled species distributions showing average range reductions of 34–40% by 2050 under various climate scenarios.^[79] For genera like Colobus, this may render up to 10% of current suitable sites uninhabitable by mid-century, compounding habitat fragmentation.^[80] Disease transmission, particularly zoonotic pathogens from increasing human proximity, adds another layer of risk, with outbreaks like Ebola devastating African primate populations. African colobines have been impacted as victims rather than reservoirs, suffering high mortality from Ebola virus disease during epidemics in Central Africa, which has led to localized extinctions in affected forests.^[81]

Protection Measures

According to assessments by the International Union for Conservation of Nature (IUCN), a high proportion of Colobinae species face extinction risks, with 96% of the 24 African colobine species classified as threatened (Vulnerable, Endangered, or Critically Endangered) and over 85% of the 28 Asian langur (Presbytis spp.) taxa similarly threatened.^[35]^[82] At least 10 species are listed as Critically Endangered, including Miss Waldron's red colobus (Piliocolobus waldroni), which is possibly extinct due to habitat loss and hunting in Ghana and Côte d'Ivoire. Key protected areas play a vital role in safeguarding Colobinae populations. In Sumatra, Indonesia, Gunung Leuser National Park harbors several endemic colobines, such as Thomas's langur (Presbytis thomasi) and the Sumatran surili (Presbytis melalophos), providing large tracts of primary rainforest essential for their folivorous diets and arboreal lifestyles. Similarly, the Dzanga-Sangha Special Reserve in the Central African Republic protects African colobines like the black-and-white colobus (Colobus guereza) and red colobus (Piliocolobus spp.), where dense lowland forests support multimale-multifemale groups amid ongoing anti-poaching efforts.^[35]^[83] International agreements further bolster protections through trade regulations and recovery initiatives. Most Colobinae species are listed under Appendix I or II of the Convention on International Trade in Endangered Species (CITES), restricting commercial trade to prevent exploitation for bushmeat, pets, or medicinal uses; for instance, the red-shanked douc langur (Pygathrix nemaeus) falls under Appendix I, prohibiting international commercial trade.^[84] Reintroduction and translocation programs have shown promise, such as the successful relocation of 17 Angolan black-and-white colobus (Colobus angolensis palliatus) in Kenya's Karura Forest, achieving a 94% survival rate and aiding habitat connectivity.^[85] Community-based conservation efforts have also contributed to recovery in some regions. In India, initiatives involving local communities around the Manas Biosphere Reserve have enhanced protection for the golden langur (Trachypithecus geei) through forest patrolling and reforestation, leading to population stabilization and increases in this Endangered species; ecotourism in the reserve generates revenue that supports anti-poaching and habitat restoration, fostering long-term stewardship.^[86]

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Nov 18, 2020 · The four species groups diverged from each other 4.5–3.1 million years ago (Ma), while speciation events within these groups occurred much more ...Mitogenomic Data · Discussion · Systematic BiologyMissing: monophyletic | Show results with:monophyletic
[15]
Adaptations to a cold climate promoted social evolution in Asian ...
Jun 2, 2023 · We constructed a socioecological-genomic framework to identify selective pressures that form the genetic basis for social evolution in Asian colobines.
[16]
[PDF] Feeding ecology of the last European colobine monkey ... - HAL
Jul 22, 2024 · If the dietary ecology of early colobines appears to be more opportunistic, it may indicate that specialized adaptations towards folivorous ...
[17]
Craniodental Mechanics and Diet in Asian Colobines - ResearchGate
May 12, 2025 · Folivory has been accepted as the general dietary pattern for colobines. However, recent ecological studies have revealed that extensive ...
[18]
Colobinae - an overview | ScienceDirect Topics
Several studies have investigated tongue morphology in the Subfamily Cercopithecinae ... Molecular analyses place the divergence time at over 60 million years.
[19]
The status ofColobus polykomos dollmani Schwarz
A supposedly intermediate subspecies,C. polykomos dollmani, is actually a hybrid swarm, in whichC. vellerosus genes greatly predominate. We propose a hypothesis ...
[20]
Old World Monkey Hybrids - Mammalian Hybrids - Macroevolution.net
Colobus guereza [Guereza Colobus] × Colobus polykomos [Western Pied Colobus] CHR. DRS. HPF. International Zoo Yearbook 1988 (p. 457); Lang 1973. Colobus ...
[21]
Taxonomic Classification of Colobine Monkeys (Chapter 2)
Feb 8, 2022 · With ten genera, 78 species and 124 taxa (species and subspecies) currently recognized, colobines are one of the most diverse primate ...
[22]
Is Malaysia's “mystery monkey” a hybrid between Nasalis larvatus ...
Apr 26, 2022 · We present evidence for a possible hybridization event between Nasalis larvatus and Trachypithecus cristatus in the Lower Kinabatangan Wildlife Sanctuary, ...
[23]
Mixed-Species Groups and Genetically Confirmed Hybridization ...
May 24, 2025 · Interspecific hybridization occurs almost in all major primate radiations. Intermediate phenotypes, sometimes seen in mixed-species groups, ...Missing: zoos | Show results with:zoos
[24]
Nuclear versus mitochondrial DNA: evidence for hybridization in ...
Colobine monkeys constitute a diverse group of primates with major radiations in Africa and Asia. However, phylogenetic relationships among genera are under ...
[25]
Sexual size dimorphism in Asian colobines revisited - PubMed
Asian colobines exhibit a wide range of sexual dimorphism in body mass. Some species are monomorphic, whereas others are strongly dimorphic.
[26]
Old World Monkeys I (Colobinae) - Encyclopedia.com
Groups show conspicuous territorial behavior. Males migrate from the natal group on reaching maturity. feeding ecology and diet. Primarily eats leaves, but ...
[27]
Colobinae - an overview | ScienceDirect Topics
Genus Cercopithecus, the vervets or guenons, is one of the most diversified genera of all primates (Figures 4.31 and 4.32). At this point in time, 26 species ...
[28]
(PDF) The Digestive Physiology of Colobine Primates - ResearchGate
Colobines are a group of primates from southeast Asia and Africa that feed, in nature, exclusively on leaves, fruits and seeds of trees.Missing: enlarged | Show results with:enlarged
[29]
https://www.researchgate.net/publication/50913863_The_Digestive_Physiology_of_Colobine_Primates
[30]
Genomic and phenotypic evidence support visual and olfactory ...
Feb 28, 2025 · ... The highest rate of olfactory receptor gene loss has been observed in colobines (Colobinae), which could be linked to the dietary transition ...Missing: capabilities | Show results with:capabilities
[31]
Cerebral complexity preceded enlarged brain size and reduced ...
Jul 3, 2015 · Such evidence indicated that the last common ancestor of cercopithecoids and hominoids had a small olfactory bulb and enhanced visual system, ...Missing: capabilities | Show results with:capabilities
[32]
(PDF) The Hand of Cercopithecoides williamsi (Mammalia, Primates)
Aug 6, 2025 · Thumb reduction is among the most important features distinguishing the African and Asian colobines from each other and from other Old World ...Missing: quadrupedalism | Show results with:quadrupedalism
[33]
None
### Summary of African Colobinae Species Distribution
[34]
Conservation of Africa's Colobine Monkeys (Cercopithecidae ...
Feb 8, 2022 · Of the monkeys in Africa, the colobines comprise 19% of the 16 genera and 30% of the 79 species. They occur all across tropical African from ...
[35]
(PDF) A taxonomic revision of the langurs and leaf monkeys (Primates
Aug 9, 2025 · Morphological and molecular systematics of the extant Colobinae Blyth, 1863 (Primates: Cercopithecidae). Article. Full-text available. Dec 2024 ...
[36]
Future simulated landscape predicts habitat loss for the Golden ...
Jun 20, 2022 · The results indicate that out of the total range extent (66,320 km2), only 12,265 km2 (18.49%) is suitable for the species at present, which ...
[37]
[PDF] Impending extinction crisis of the world's primates
Jan 18, 2017 · A rapid expansion of agriculture in primate range regions has been paralleled by a sharp decline in forest cover in the 20-year period ...
[38]
(PDF) Behavior, Ecology, and Conservation of Colobine Monkeys
Aug 9, 2025 · Colobine monkeys are distributed in sub-Saharan African forests and all over the South and Southeast Asia (Fashing 2007) . The extant colobine ...
[39]
Population structure and distribution of geladas (Theropithicus ...
Jun 28, 2023 · The area comprises five dominant habitat types namely; grassland, wooded grassland, natural forest (dry evergreen Afromontane forest), ...
[40]
The importance of well protected forests for the conservation ...
Both species are also found in the largest and best‐preserved West African forest—the Taï National Park (TNP) in Ivory Coast. Colobine monkeys are hunted for ...
[41]
Seasonal variation in activity budgets of critically endangered ...
May 15, 2025 · The langurs were observed more frequently during the dry season (345 min) compared to the wet season (235 min).
[42]
A multi-level society comprised of one-male and multi-male core ...
Oct 16, 2019 · Abstract. Several mammalian species exhibit complex, nested social organizations, termed multi-level or modular societies.Missing: Colobinae | Show results with:Colobinae
[43]
Asian colobine social structure: Ecological and evolutionary ...
As a group, the Asian colobines may be partially buffered from the impact of food constraints on social structure, given their specialized digestive physiology.
[44]
Socioecology of African Colobines (Chapter 17)
Feb 8, 2022 · These dominance hierarchies tend to be individualistic and females typically do not stay with kin to defend food. Unfortunately, there are still ...
[45]
Dominance hierarchy and social relationships in a group of Captive ...
A strong dominance hierarchy is characterized by common agonistic1interactions, and less than 5% of reversals indicating stable dominance relationships. By ...
[46]
Within-and among-male variation in roaring by black and white ...
Aug 9, 2025 · This study tests whether some male black and white colobus monkeys (Colobus guereza, 'guerezas') indicate their willingness to defend access to ...
[47]
Primate Sociality and Social Systems | Learn Science at Scitable
One of the most common primate social systems is the one-male group, which characterizes most colobine monkeys, most guenons, patas monkeys, howler monkeys, and ...<|control11|><|separator|>
[48]
The Evolution of Allomothering Behavior Among Colobine Monkeys ...
The great majority of Old World monkey females are interested in infants, particularly very young ones. Characteristically, females will approach infants ...
[49]
Costs and benefits of allomaternal care to mothers and others in wild ...
Oct 1, 2024 · Evolution of allomothering behavior among colobine monkeys: Function and opportunism in evolution. American Anthropologist, 81, 818–840 ...Missing: Colobinae | Show results with:Colobinae<|separator|>
[50]
Causes and Consequences of the Formation of Multilevel Societies ...
Feb 8, 2022 · Colobines display an array of different types of social organization. In some Asian colobines (most prominently the snub-nosed monkeys, ...
[51]
Diet and Activity Budget in Colobus angolensis ruwenzorii at ...
Oct 30, 2020 · Colobine monkeys are specialized folivores that use foregut fermentation to digest leaves. The slow process of fermentation forces them to spend ...<|control11|><|separator|>
[52]
[PDF] Foraging challenges of red colobus monkeys - Squarespace
However, when primates eat foliage they are very selective of species and plant parts, feeding primarily on young leaves of only a few species. Protein and ...
[53]
Colobine Nutritional Ecology (Chapter 8) - Cambridge University Press
Feb 8, 2022 · It follows that colobines prefer mature and young leaves with higher protein-to-fibre ratios compared to those leaves less preferred or uneaten.
[54]
Foraging challenges of red colobus monkeys - PubMed
Previous studies have found the biomass of folivorous primates to be related to the ratio of protein to fiber concentration of mature leaves in the environment.Missing: scientific | Show results with:scientific
[55]
Natural History of Black-and-White Colobus Monkeys (Chapter 10)
Feb 8, 2022 · The black-and-white colobus monkeys consist of five arboreal species classified in the genus Colobus (C. angolensis, C. guereza, C. polykomos, C ...<|control11|><|separator|>
[56]
Ranging and foraging of Himalayan grey langurs (Semnopithecus ...
Feb 14, 2013 · The daily path lengths of the SBG, LBG, and AMG were 1.23 ± 0.28 (mean ± SD; average of four seasons), 1.75 ± 0.34, and 1.84 ± 0.70 km, ...Missing: distance | Show results with:distance
[57]
Seasonal variation in activity budgets of critically endangered ...
The langurs predominantly use stratum C (21–30 m) to feed, rest, move, and socialize. This study highlights the seasonal ecology of this critically endangered ...Missing: migration | Show results with:migration
[58]
[PDF] Anti-predator behaviour of Guereza colobus monkeys
Predator abundance, predation risks, predator hunting techniques, Guereza anti-predator ... predator's biological class differed between the populations ...
[59]
[PDF] Behaviours of wild male and female Colobus angolensis palliatus ...
a palliatus strategy to avoid predators. Colobines balance their activities for several reasons and predator avoidance is one of them (Stanford, 2001). The ...
[60]
Observations on the Sexual Behaviour and Birth Seasonality of ...
monkey reproductive success, with a polygynous mating system, adult male proboscis monkeys marry many adult female proboscis monkeys (Boonratana, 2011 ).
[61]
Evolutionary determinants of modular societies in colobines
Modular societies are structurally characterized by nuclear one-male units (OMUs, or harems) embedded within larger relatively coherent social bands.
[62]
Socioecology of Asian Colobines (Chapter 16)
Feb 8, 2022 · In proboscis monkeys (Nasalis larvatus) aggression is rare, yet females have conflicts for sleeping sites and, while quantitative data are not ...
[63]
Sexually selected lip colour indicates male group-holding status in ...
Dec 16, 2015 · Snub-nosed monkeys have a one-male unit (OMU) based ... The mating system of the Sichuan snub-nosed monkey (Rhinopithecus roxellana).
[64]
Behavioral and endocrine dynamics associated with infanticide in a ...
Infanticide has been observed in several colobines, but only one infanticide has previously been documented for black and white colobus monkeys (Colobus guereza) ...Missing: systems polygyny
[65]
[PDF] 16 Socioecology of Asian Colobines - UU Research Portal
Therefore, the lack of nepotism in the colobine species with linear female dominance hierarchies and where female relatives are available, such as some well- ...
[66]
toward an understanding of their life history and mating system
To learn more about the life history characteristics and mating system of guerezas, we documented the mating behavior and ovarian hormone profiles of 10 cycling ...Missing: Colobinae review
[67]
Red colobus monkeys display alternative behavioral responses to ...
Controlling for ecological variation in resource availability is critical ... interbirth intervals vary widely with an average of about 23 months ...
[68]
Ecology of Semnopithecus (Chapter 13) - The Colobines
Feb 8, 2022 · Early work by Hladik (Reference Hladik and Clutton-Brock1977) found that purple-faced langurs exploited more leaves and less fruit than ...Missing: migration fruiting cycles
[69]
Infanticide as Sexual Conflict: Coevolution of Male Strategies and ...
The original sexual selection hypothesis arguing that infanticide improves male mating success by accelerating the return of females to fertilizable ...Missing: Colobinae | Show results with:Colobinae
[70]
Table of Primate Reproduction Data
The below table is extracted from the old PIN and includes mating system, age at first birth, gestation length, litter size, birth rate, neonate weight, age ...
[71]
Mantled guereza (colobus)
TAXONOMY. Suborder: Haplorrhini ... Variation in the diets of Cercopithecus species: differences within forests, among forests, and across species.
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Primate life histories and dietary adaptations: A comparison of Asian ...
Oct 5, 2010 · The current analysis found a longer gestation period for colobines, a similar age at first birth, and a similar interbirth interval, but did not ...
[73]
Weaning Age, Infant Care, and Behavioral Development in ...
May 17, 2008 · Weaning Age, Infant Care, and Behavioral Development in Trachypithecus leucocephalus. Published: 17 May 2008. Volume 29, pages 583–591, (2008) ...
[74]
Maternal care and infant transfer in two species ofColobus monkeys
Aug 5, 2025 · During early development, non-mothers performed more maternal behaviors and transferred and carried the infant more than its mother. With infant ...
[75]
Colobus angolensis - Animal Diversity Web
Females of the family group mate with the dominant male. Mating System; polygynous polygynous. In most Angolan colobus social groups there is one adult male ...
[76]
Piliocolobus rufomitratus (Tana river red colobus) | INFORMATION
In general, red colobus monkeys breed throughout the year. Estimated gestation is 4.5 to 5.5 months, with an estimated interbirth interval of 26 months. The ...
[77]
Red-Shanked Douc Langur, Pygathrix nemaeus
The International Union for Conservation of Nature lists red-shanked douc langurs as Critically Endangered (IUCN, 2015), appearing on the IUCN Red List of ...
[78]
Allen's swamp monkey - National Zoo
The primary threats to the monkeys include hunting for bushmeat, killing in retaliation for crop raids and collection for the pet trade. IUCN states a need ...Missing: Colobinae | Show results with:Colobinae
[79]
Over 9,000 primates killed for single bushmeat market in West Africa ...
Mar 24, 2014 · Over the past 25 years, West Africa's primates have been put at risk due to an escalating bushmeat trade compounded with forest loss from expanding human ...Missing: Allen's swamp<|separator|>
[80]
Climate change projections suggest severe decreases in the ...
We found that by 2050, under MIROC5, modeled species distributions showed an average reduction of 34%, and under CCSM4, a decrease of 40%. By 2070, MIROC5 ...Missing: Colobinae | Show results with:Colobinae
[81]
The Effect of Climate Change on the Distribution of the Genera ...
The Colobus genera will see a marked decrease of climatically suitable areas, whereas projections are more contrasted for Cercopithecus species, which will also ...
[82]
African Primates: Likely Victims, Not Reservoirs, of Ebolaviruses - PMC
Extensive serological survey of multiple African non-human primate species reveals low prevalence of IgG antibodies to four Ebola virus species.Missing: colobines | Show results with:colobines
[83]
[PDF] Asian langurs (Presbytis) conservation action plan 2024–2034
According to the IUCN Red List of Threatened SpeciesTM, 24 of the. 28 taxa (>85%) are threatened with extinction (listed as. Critically Endangered, Endangered ...
[84]
Facts and Infos – Dzanga-Sangha Protected Area
Part of the “Northwestern Congolian lowland Forests”, Dzanga-Sangha Protected Areas are located in the southwestern corner of the Central African Republic, ...Missing: colobines | Show results with:colobines
[85]
Appendices | CITES
In addition, all parts and derivatives thereof are also included in the same Appendix unless, for animal species listed in Appendix III and plant species listed ...<|control11|><|separator|>
[86]
Colobus Project Success – Friends of Karura Forest
The Karura Colobus monkey project is one of the most successful translocation ever undertaken for arboreal primates with a success rate of 94%!Missing: langur | Show results with:langur
[87]
Community protection of the Manas Biosphere Reserve in Assam ...
Apr 6, 2010 · The Golden Langur Conservation Project has resulted in an increase in the total Indian population of golden langurs, control of illegal logging ...Missing: stabilization | Show results with:stabilization