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Semnopithecus

Semnopithecus is a genus of Old World monkeys in the family Cercopithecidae, subfamily Colobinae, commonly known as gray langurs or Hanuman langurs, characterized by their predominantly folivorous diet, arboreal lifestyle, and adaptability to diverse habitats across South Asia.^[1] The genus Semnopithecus is classified within the tribe Presbytini and includes eight species: S. schistaceus (Nepal gray langur), S. ajax (Kashmir gray langur), S. hector (Tarai gray langur), S. entellus (northern plains gray langur), S. hypoleucos (black-footed gray langur), S. priam (tufted gray langur), S. johnii (Nilgiri langur), and S. vetulus (purple-faced langur).^[2] These species exhibit varying morphologies, with some debate in taxonomy due to hybridization and clinal variation, but molecular and morphological studies support this division.^[3] Gray langurs are medium to large-sized primates, with males weighing 14–24 kg and females 10–15 kg, featuring silvery-gray fur, a black face, and a long tail often exceeding body length for balance during arboreal locomotion.^[1] They inhabit a wide range of environments, from tropical rainforests and deciduous forests in the Indian subcontinent to temperate Himalayan forests, grasslands, and even urban areas in Pakistan, India, Nepal, Bhutan, Bangladesh, and Sri Lanka.^[1]^[4] Socially, Semnopithecus species live in flexible groups of 10–100 individuals, often structured as one-male units with all-male bands or multimale-multifemale troops, where males may engage in infanticide to bring females into estrus.^[1] Their diet is primarily leaves, supplemented by fruits, seeds, and flowers, supported by specialized sacculated stomachs for fermenting fibrous vegetation.^[1] Reproduction is seasonal in some populations, with females giving birth to single offspring after a 200-day gestation, and groups showing cooperative care.^[4] While adaptable, several species face threats from habitat loss and human-wildlife conflict, leading to varying conservation statuses from least concern to endangered.^[4]

Physical description

Morphology

Species of the genus Semnopithecus, commonly known as gray langurs, exhibit moderate body sizes typical of arboreal colobine monkeys. Adult head-body lengths range from 50 to 70 cm, with tails measuring 70 to 100 cm, often exceeding the body length to aid in navigation through forested environments.^[5]^[4] Body weights vary by species and sex, generally falling between 9 and 20 kg, with males typically larger than females; for instance, in S. entellus, males average 13 kg and females 9.9 kg.^[6]^[7] Their limb structure is adapted for life in arboreal and semi-terrestrial habitats, featuring elongated fore- and hindlimbs that facilitate quadrupedal walking, climbing, and leaping between branches.^[1] These long limbs enable efficient traversal of discontinuous supports in trees, while the hands possess opposable thumbs, allowing precise grasping of leaves and branches during foraging.^[8] The overall skeletal morphology supports both terrestrial quadrupedalism and occasional suspension, reflecting their opportunistic use of ground and canopy spaces.^[1] Facial features include a pronounced muzzle suited to their folivorous diet and large, forward-facing eyes that enhance binocular vision for diurnal activity.^[9] A key anatomical adaptation is the specialized sacculated stomach, characteristic of colobine primates, which consists of multiple chambers—the rumen-like forestomach for microbial fermentation of fibrous plant material, followed by chambers for further digestion and absorption.^[10] This foregut fermentation system, supported by symbiotic bacteria, allows efficient breakdown of leaves and other tough vegetation, enabling Semnopithecus species to thrive on a diet high in cellulose.^[11] The tail serves primarily as a counterbalance during locomotion, stabilizing the body during leaps and climbs in arboreal settings.^[12] Additionally, it functions in social communication, where movements such as waving or positioning can convey signals related to dominance, submission, or affiliation within groups.

Coloration and sexual dimorphism

Semnopithecus species possess a dense, long, and silky fur coat that provides insulation and protection in their varied habitats. The typical coloration is predominantly gray, often with a silvery or brownish tinge on the dorsal surface and lighter, yellowish-white fur on the ventral side, while faces, hands, and feet are usually black. This pelage varies subtly across individuals.^[4]^[6] Coloration differs markedly among species, reflecting adaptations to local environments. For instance, the Nilgiri langur (S. johnii) features glossy black fur across the body with a distinctive golden-brown mane on the head, enhancing its contrast against forested backdrops. In contrast, the tufted gray langur (S. priam) displays gray to brownish-gray dorsal fur that darkens with age, paler underparts, and a prominent upward-pointing tuft of crown hair, with black facial skin. The northern plains gray langur (S. entellus) shows light gray overall with darker shoulders and tail base, incorporating reddish tinges on the back.^[13]^[14]^[15]^[4] Sexual dimorphism in Semnopithecus is primarily manifested in body size, with males generally larger and more robust than females, though pelage differences are minimal. Males often exhibit more pronounced facial markings or crests, such as enhanced tufts in S. priam, while females are slightly smaller with subdued color intensity. A notable exception occurs in S. johnii, where females possess unique white patches on the inner thighs absent in males, potentially aiding in infant visibility or thermoregulation. Beyond these traits, no substantial differences in fur coloration or texture exist between sexes.^[5]^[13]^[15] The gray tones prevalent in Semnopithecus fur serve a critical role in camouflage, blending seamlessly with the muted hues of forest canopies, rocky terrains, and arid scrublands to evade predators. This countershading—darker dorsally and lighter ventrally—further aids concealment by mimicking shadows and light patterns in their arboreal and terrestrial environments.^[16]^[4]

Taxonomy and phylogeny

Species and subspecies

The genus Semnopithecus currently includes eight recognized species as of 2022, reflecting recent taxonomic revisions based on morphological, genetic, and distributional data.^[17] These species are distinguished primarily by variations in pelage color, limb darkness, crest morphology, and tail characteristics, which aid in field identification.^[4] The northern plains gray langur (S. entellus) is characterized by its pale gray dorsal fur, lighter ventral pelage, and a uniformly colored tail without distinct banding; it includes several subspecies, such as S. e. entellus in central India, which features a more pronounced occipital crest compared to northern variants.^[4] The black-footed gray langur (S. hypoleucos) differs with darker hands and feet contrasting against its overall gray body and a slightly darker crown. The tufted gray langur (S. priam) exhibits a distinctive tuft of hair on the crown and darker overall coloration, with subspecies like S. p. thersites showing even more pronounced facial markings in southern populations.^[4] The Nilgiri langur (S. johnii) is notable for its blackish pelage, white ventrum, and short tail, adapted to high-altitude forests.^[18] The Nepal gray langur (S. schistaceus), elevated to full species status in 2020 based on genetic analyses confirming divergence from S. entellus, features a darker slate-gray coat, robust build, and a flat crown without tufts. The Tarai gray langur (S. hector) has a yellowish-gray hue with brown limbs and a pale belly, while the southern plains gray langur (S. dussumieri) shows a reddish tinge to its gray fur and darker extremities. Finally, the Kashmir gray langur (S. ajax) displays a silvery-gray pelage with blackish limbs and a long, thin tail.^[4]^[19] Hybridization has been observed between closely related species in zones of sympatry, such as between S. priam and S. vetulus (sometimes classified within Semnopithecus), indicating ongoing gene flow that complicates taxonomic boundaries.^[20]

Classification history

Semnopithecus belongs to the subfamily Colobinae within the family Cercopithecidae, comprising Asian leaf monkeys and langurs. Molecular phylogenetic analyses place Semnopithecus as sister to Trachypithecus, with Presbytis sister to this pair and the odd-nosed colobines (e.g., Rhinopithecus) among the Asian colobines.^[21] This positioning is supported by both mitochondrial and nuclear DNA data, indicating that the common ancestor of Semnopithecus, Trachypithecus, and Presbytis diverged from other Asian colobine lineages approximately 10–8 million years ago during the late Miocene, based on relaxed molecular clock estimates calibrated with fossil constraints.^[22] Historically, the genus Semnopithecus was considered monotypic, encompassing only S. entellus (the northern plains gray langur), with regional variants treated as subspecies based primarily on morphological traits such as pelage color and tail length.^[23] This view persisted until the early 2000s, when genetic studies began revealing deeper divergences within the group. A pivotal 2008 molecular analysis using complete mitochondrial genomes clarified the phylogenetic distinction between Semnopithecus and Trachypithecus, supporting the recognition of multiple species within Semnopithecus and restricting the genus to the Indian subcontinent.^[24] These findings prompted taxonomic revisions, shifting from a single polytypic species to a multispecies framework driven by evidence of genetic isolation corresponding to geographic barriers. Key taxonomic revisions include the 2004 proposal by Brandon-Jones, which reduced the number of subspecies under S. entellus to seven and elevated the southern populations to the species S. priam based on morphometric and distributional data.^[25] More recently, a 2020 integrative study using morphology, genetics, and ecology confirmed the species status of the Himalayan gray langur as S. schistaceus, distinguishing it from S. entellus through fixed cranial differences and mitochondrial divergence.^[19] The placement of the purple-faced langur (S. vetulus) remains debated; while some genetic evidence supports inclusion in Semnopithecus, it is often treated as a distinct lineage or reassigned to Trachypithecus due to phylogenetic incongruences.^[26] The fossil record of Semnopithecus is limited, with no definitive Miocene representatives, but it is linked to early colobine radiations in Asia. The genus likely evolved from Miocene ancestors like Mesopithecus, known from late Miocene sites across Eurasia, including western and eastern Asia, around 10–8 million years ago.^[27] The earliest attributed Semnopithecus fossils date to the late Pliocene (approximately 3.5 million years ago) from central Myanmar, consisting of dental and mandibular remains that exhibit derived folivorous adaptations similar to extant species.

Distribution and habitats

Geographic range

The genus Semnopithecus is primarily distributed across the Indian subcontinent, encompassing India, Sri Lanka, Nepal, and Bangladesh, with some species extending into Pakistan and Bhutan.^[1] This range reflects the genus's adaptation to diverse environments, though distributions vary significantly by species.^[4] Species-specific ranges highlight regional specialization. The northern plains gray langur (S. entellus) occupies a broad area in the plains of central and northern India south of the Himalayan foothills, extending to the Narmada and Krishna rivers in the south, and is also present in Bangladesh.^[28] The Nepal gray langur (S. schistaceus) has the most extensive Himalayan distribution, spanning Nepal, northern India, northern Pakistan, Bhutan, and far southwestern Tibet.^[12] In contrast, the Kashmir gray langur (S. ajax) is confined to the Himalayan tracts of northern India and Pakistan.^[4] The Tarai gray langur (S. hector) occurs in the Himalayan foothills of northern India, Nepal, and Bhutan, at altitudes of 150–1,600 m. The southern plains gray langur (S. dussumieri) ranges through west-central and southwestern India.^[4] The Nilgiri langur (S. johnii) is highly restricted to fragmented pockets in the Western Ghats, including the Nilgiri-Brahmagiri, Siruvani, Anamalai-Parambikulam, Cardamom Hills, and Periyar-Agastyamalai regions.^[29] The tufted gray langur (S. priam) occurs in southern India and Sri Lanka.^[15] Compared to historical distributions, the current range of Semnopithecus has contracted in some areas due to habitat fragmentation from human expansion, though the genus's adaptability has enabled persistence in modified landscapes.^[30] For instance, S. entellus populations have declined in rural forested zones but thrive in urban settings like Delhi through opportunistic foraging.^[30] Overlap zones exist in northern and central India, where species such as S. entellus and S. schistaceus coexist near riverine boundaries, potentially facilitating gene flow between populations.^[28]

Habitat types

Semnopithecus species exhibit remarkable adaptability across a variety of forest habitats, primarily occupying tropical dry and deciduous forests as well as moist evergreen forests throughout South Asia. These monkeys show a strong preference for the middle canopy layers, typically between 5 and 15 meters in height, where dense foliage provides optimal cover and foraging opportunities.^[1]^[4]^[31] Beyond forested environments, Semnopithecus demonstrates high tolerance for non-forest adaptations, including open woodlands, savannas, and shrublands, as well as heavily disturbed areas such as agricultural fields and urban fringes. This ecological flexibility allows populations to persist in human-modified landscapes, where they often exploit proximity to human settlements for supplemental resources.^[5]^[6] The genus occupies a broad altitudinal range, from sea level to elevations exceeding 4,000 meters, with species like Semnopithecus schistaceus thriving in Himalayan highlands up to 4,000 meters above sea level. Microhabitat preferences include sites near water sources for hydration, supplemented by moisture from foliage, and sleeping sites in tall trees or rocky cliffs for protection from predators.^[19]^[12]^[13]

Ecology

Diet and foraging

Semnopithecus species are predominantly folivorous primates, with leaves forming the bulk of their diet, typically comprising 50-85% of intake across various populations and habitats. Fruits (15-25%), flowers (4-13%), seeds, and other plant parts constitute the primary supplements, while insects (0.4-3%), bark, stems, roots, moss, soil, and occasionally cultivated crops make up minor portions. This dietary specialization enables them to exploit abundant but nutritionally poor, high-fiber foliage that is less accessible to other primates.^[4]^[32]^[7] Their digestive system is adapted for processing such low-quality foods through a complex, multi-chambered foregut featuring a sacculated stomach that supports microbial fermentation. Symbiotic bacteria in the rumen-like chambers break down cellulose and other fibers, extracting energy from otherwise indigestible plant material via volatile fatty acid production. This foregut fermentation system distinguishes colobines like Semnopithecus from most other primates and allows efficient utilization of mature leaves high in structural carbohydrates.^[33]^[7] Foraging occurs diurnally, with individuals and groups visually scanning tree canopies to select food items, prioritizing young leaves when available but shifting to mature foliage as fallback resources during periods of scarcity. Seasonal variations influence these patterns, as wetter periods increase fruit availability and consumption, while dry seasons emphasize persistent leaf resources to maintain nutritional stability. Gut microbiota further support this ecology by detoxifying tannins, alkaloids, and other plant secondary compounds prevalent in mature leaves, enabling safe ingestion of toxin-laden fallback foods.^[4]^[34]^[10] To compensate for the low energy density of their diet, Semnopithecus consume substantial daily volumes of fresh plant matter, which sustains metabolic demands through high throughput and microbial efficiency.

Activity patterns

Semnopithecus species exhibit a distinctly diurnal activity pattern, remaining active from dawn to dusk while spending nights resting or sleeping in trees to avoid nocturnal predators. Peak activity periods occur mid-morning and in the afternoon, when individuals engage in foraging, locomotion, and social interactions, with resting dominating midday hours. This cycle aligns with environmental light availability and resource accessibility, allowing efficient energy expenditure in their arboreal and terrestrial lifestyles.^[4]^[8] Seasonal variations influence activity rhythms, with reduced overall movement and increased resting during hot or dry seasons to conserve energy amid scarce resources, while monsoon periods see heightened ranging and foraging due to abundant vegetation. In Himalayan populations, such as Semnopithecus ajax, daily path lengths extend longer during winter months, averaging up to 1.84 km, compared to shorter distances in resource-richer seasons. These adaptations reflect responsiveness to climatic and phenological changes, maintaining group cohesion and nutritional needs.^[35]^[36] Ranging behavior varies by group size, habitat type, and resource distribution, with home ranges typically spanning 0.5 to 5 km² for bisexual troops, though larger all-male bands may exceed 10 km² in expansive terrains like the Himalayas. Daily path lengths generally measure 1 to 2 km, enabling coverage of feeding sites without excessive energy cost, and these patterns integrate briefly with foraging routines to optimize resource exploitation.^[37]^[38] Langurs demonstrate heightened alertness to disturbances, such as predators or human proximity, by increasing vigilance, emitting alarm calls, or performing defensive actions like direct confrontation, which temporarily disrupts normal activity flows. Nocturnal inactivity prevails in natural habitats, though urban-adapted groups may show extended activity into evenings due to provisioning and reduced diurnal threats.^[39]^[4]

Behavior

Semnopithecus species exhibit flexible social structures, primarily organized into multimale-multifemale troops consisting of 10 to 60 individuals, though group sizes vary with habitat quality and population density. These troops form the core social unit, comprising multiple adult females, their offspring, and several adult males, with females serving as the stable, kin-based foundation due to lifelong female philopatry. Males typically disperse from their natal groups at maturity, joining all-male bands or attempting to enter existing troops, which contributes to the dynamic nature of male membership. All-male bands, ranging from a few to over 30 individuals, serve as transient groups for subadult and peripheral males awaiting opportunities to integrate into bisexual troops.^[8]^[40]^[5] Within troops, dominance hierarchies structure social interactions, with males establishing rank through agonistic displays, chases, and occasional fights, often led by a central alpha male who directs group movement and defends resources. Female hierarchies are stable and linear, typically matrilineally inherited among related kin groups, influencing access to food and grooming partners without the intense aggression seen in males. This dual hierarchy promotes cohesion, as females bond through affiliative behaviors while males compete for reproductive access and tenure.^[41]^[42]^[43] Intergroup dynamics involve territorial defense, where coalitions of resident males confront intruders through vocalizations and physical clashes to protect core areas rich in resources. Peaceful associations may occur at abundant feeding sites, allowing temporary overlap without escalation. A notable aspect is male takeovers, during which incoming males from all-male bands may evict residents and commit infanticide against unrelated infants to accelerate female reproductive cycles, a strategy observed across multiple populations and linked to high male turnover rates.^[38]^[44] Groups occasionally undergo fission in response to resource scarcity or excessive size, splitting into smaller subunits that may reform or establish independently, while fusion is rare but can occur through male-mediated mergers of fragments. Female philopatry ensures the persistence of matrilineal bonds across such events, maintaining troop stability over generations.^[45]^[46]

Reproduction and parental care

Semnopithecus species exhibit a polygynandrous mating system, characterized by multimale-multifemale troops where females mate with multiple males, often preferentially with the alpha male, though subordinate males also sire offspring. Mating activity typically peaks during the dry season, aligning with resource availability and female estrus cycles that last approximately 24-26 days.^[4]^[47]^[48] Reproductive biology in Semnopithecus involves a gestation period of 190-210 days, usually resulting in a single offspring, though twins occur rarely. Females reach sexual maturity between 3 and 4 years of age, while males mature at 5-6 years, after which females remain in their natal group and males disperse to form all-male bands or challenge for troop residency. Births predominantly occur at night, with infants initially clinging ventrally to the mother before shifting to dorsal carriage after 1-2 weeks.^[49]^[4]^[50] Parental care is primarily maternal but extensively supported by allomothering, where non-mother females and subadults groom, carry, and even nurse infants, particularly during the first month postpartum, which helps reduce maternal energetic costs and enhances infant survival. Adult males provide protection against predators and conspecific threats, tolerating and sometimes grooming young to build social bonds, though this can shift during takeovers. Weaning typically begins around 8-10 months and completes by 12-13 months, after which juveniles integrate more fully into group foraging. Infant mortality is notably high, often exceeding 30% in the first year, largely due to infanticide perpetrated by incoming males aiming to accelerate female fertility.^[51]^[4]^[47] Life history traits reflect the genus's adaptation to variable environments, with an interbirth interval of 2-3 years for successfully reared offspring, influenced by infant survival and resource conditions. In the wild, longevity averages 20-30 years, with females outliving males due to lower risks from intermale aggression and dispersal.^[48]^[6]^[52]

Communication

Semnopithecus species exhibit a diverse vocal repertoire comprising at least 19 distinct call types, serving functions in social coordination, aggression, and predator defense.^[5] Adult males frequently produce loud whoops or calls to signal group movements, finalize decisions, and during intergroup displays, while all group members use grunts, screams, and barks in various contexts such as distress or fights.^[53]^[4] Alarm calls, including harsh barks and short barks, are emitted by adults and subadults upon detecting predators, with loud barks specifically directed at ground threats like leopards to alert the group and nearby species.^[54] These vocalizations often elicit rapid anti-predator responses, such as mobbing or fleeing, enhancing group survival.^[5] Visual signals play a key role in intra- and intergroup interactions among Semnopithecus, including threat displays like staring, open-mouth grimaces, and mounting behaviors to assert dominance within the social hierarchy.^[4] Females employ head shaking as a visual cue to indicate estrous status, facilitating mate attraction.^[6] Mothers use infant-presenting gestures, holding juveniles toward other group members, to solicit grooming or reduce aggression during tense encounters.^[4] In intergroup challenges, males perform branch-shaking displays accompanied by vocalizations to deter rivals without physical contact.^[55] Tactile communication is essential for maintaining social bonds and resolving conflicts in Semnopithecus groups, with allogrooming serving as a primary mechanism to reinforce alliances and reduce tension.^[6] Dominant individuals both give and receive grooming more frequently, strengthening their position in the hierarchy, while post-conflict embraces and clasping behaviors aid in reconciliation.^[56]^[57] Females may emit specific copulation calls during mating to signal satisfaction or influence male behavior, aligning with broader patterns in colobine primates.^[58] These multimodal signals collectively facilitate complex intra- and intergroup dynamics, from predator avoidance to reproductive interactions.

Conservation

Threats

Semnopithecus populations face significant threats from habitat loss primarily driven by deforestation for agriculture, urbanization, and infrastructure development, which fragments forests and reduces available foliage essential for their folivorous diet. In species like the northern plains gray langur (S. entellus), intensive agriculture and expanding human settlements have led to widespread habitat degradation across much of their range in northern India and Bangladesh.^[59] Similarly, in the Western Ghats, the Nilgiri langur (S. johnii) experiences high impacts from habitat fragmentation due to tea plantations and logging, contributing to its Vulnerable status.^[60] Overall, habitat loss affects approximately 76% of primate species, including Semnopithecus, underscoring the scale of anthropogenic pressure on their distributions.^[30] Poaching and human persecution further endanger these langurs, with individuals hunted for bushmeat or killed in retaliation for crop raiding in agricultural areas. The Nilgiri langur is particularly vulnerable to poaching for its pelt, organs, blood, and flesh, believed to have medicinal value in local traditions.^[61] Human-wildlife conflict exacerbates this, as langurs foraging near farmlands are often targeted, while roadkill from vehicle collisions in expanding urban fringes poses an additional mortality risk, as documented in regions like Rajasthan, India.^[4] Natural threats include predation by large carnivores such as leopards (Panthera pardus) and tigers (P. tigris), which preferentially target langurs in forested habitats, especially during periods of low prey availability.^[4] Disease transmission from humans and domestic animals also poses risks, with rabies spillover from dogs reported in cases involving gray langurs, leading to fatal infections in affected individuals.^[62]^[63] Climate change compounds these pressures by altering monsoon patterns, which disrupts leaf availability and foraging resources in tropical and subtropical habitats. For the Kashmir gray langur (S. ajax), projected range shifts toward higher elevations are expected by 2050 due to warming temperatures and changing precipitation, potentially reducing suitable habitat by over 60% under various scenarios.^[64]^[65]

Protection efforts

The conservation status of Semnopithecus species varies across the genus (as assessed by the IUCN in 2015–2020), with several classified as Vulnerable or Endangered due to habitat fragmentation and localized declines, while others remain Least Concern. For instance, the Nilgiri langur (S. johnii) is listed as Vulnerable, reflecting ongoing population reductions in its restricted range in the Western Ghats (2015). The Kashmir gray langur (S. ajax) is assessed as Endangered (2015), primarily owing to habitat loss and human pressures in its high-altitude habitats. The Tarai gray langur (S. hector) is Near Threatened (2015). In contrast, the northern plains gray langur (S. entellus) and Nepal gray langur (S. schistaceus) are categorized as Least Concern (2015), benefiting from wider distributions and cultural protections in parts of their range. The tufted gray langur (S. priam) holds a Near Threatened status (2015), highlighting moderate risks from agricultural expansion. The southern plains gray langur (S. dussumieri) is Least Concern (2015).^[66] Legal frameworks provide critical safeguards for Semnopithecus across their range. In India, most species fall under Schedule II of the Wildlife (Protection) Act, 1972, prohibiting hunting, trade, and capture with penalties for violations, though enforcement challenges persist. The Nilgiri langur receives heightened protection under Schedule I, Part I, offering stricter prohibitions and higher penalties due to its endangered status. Several species, including S. entellus and S. ajax, are listed in Appendix I of the Convention on International Trade in Endangered Species (CITES), banning international commercial trade to prevent exploitation.^[67] In Sri Lanka, the tufted gray langur (S. priam) is protected under the Fauna and Flora Protection Ordinance, which regulates habitat disturbance and poaching through national park designations and licensing requirements. Conservation initiatives emphasize habitat preservation and human-wildlife coexistence. Protected areas such as Silent Valley National Park in India safeguard the Nilgiri langur through core forest reserves, while reforestation projects in the Western Ghats, including community-led planting of native trees, aim to reconnect fragmented habitats for species like S. johnii, enhancing corridor connectivity.^[68] Community-based anti-poaching efforts, such as village patrols in the Himalayas, have reduced illegal hunting of the Kashmir gray langur by involving local stakeholders in monitoring and awareness campaigns. Ongoing research and monitoring support targeted interventions. Genetic studies using mitochondrial DNA have clarified subspecies boundaries for S. johnii and S. ajax, informing prioritization of distinct populations for protection.^[60] Camera trap surveys provide population estimates for elusive groups, such as in fragmented forests. Ecotourism programs in urban-adjacent areas, like those near temple sites in India, promote tolerance of S. entellus troops by generating revenue for conservation while educating visitors on their ecological role.^[4]

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Distribution, social organization, and management of ...
Nov 29, 2023 · Nilgiri langurs are found in five regions in the Western Ghats: Nilgiri-Brahmagiri, Siruvani, Anamalai- Parambikulam, Cardamom Hills, and Periyar-Agastyamalai.
[28]
Impending extinction crisis of the world's primates - PubMed Central
Baboons (Papio), Hanuman langurs (Semnopithecus), and macaques (Macaca) are particularly adaptable and can survive even in urban areas (49). Chimpanzees appear ...
[29]
Activity budget and Communication Dynamics of Black leaf monkeys ...
Sep 21, 2025 · They preferred the middle canopy layer (>5-15m). Time-activity analysis revealed that resting (61%) and eating (23%) dominated their daily ...
[30]
https://pmc.ncbi.nlm.nih.gov/articles/PMC5242557/
[31]
Morphology and Physiology of Colobine Digestive Tracts (Chapter 6)
Feb 8, 2022 · Colobine monkeys differ from all other primates in having a foregut fermentation system featuring a complex, multi-chambered stomach.
[32]
Optimal foraging on the roof of the world: Himalayan langurs and the ...
Oct 20, 2009 · These results suggest that even simple models from foraging theory can provide a useful framework for the study of primate feeding behavior.
[33]
Specialized digestive adaptations within the hindgut of a colobine ...
Mar 29, 2022 · GI microbes also neutralize dietary toxins and digestive inhibitors,, further improving host digestive efficiency. These functions are ...Missing: avoidance mature
[34]
131 Purple-faced Langur (Semnopithecus vetulus) also ... - Facebook
Jan 28, 2019 · It has a dark brown to black coat, a distinctive purplish face with white fur around the mouth, and a long tail for balance.Missing: shedding | Show results with:shedding
[35]
Ranging and foraging of Himalayan grey langurs (Semnopithecus ...
Feb 14, 2013 · We conducted a 1-year study of Himalayan grey langurs (Semnopithecus ajax) in Machiara National Park, Pakistan, where they occur at relatively high altitudes.Missing: altitudinal | Show results with:altitudinal
[36]
Ranging and foraging of Himalayan grey langurs (Semnopithecus ...
Aug 6, 2025 · 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, ...<|separator|>
[37]
Ranging Behaviour of Hanuman Langurs (Semnopithecus entellus ...
Aug 1, 2006 · The home ranges were 106, 45, and 70 ha. This study suggests that the availability of preferred foods, including farm crops and provisioned food ...
[38]
Anti-predator strategies of adult male Central Himalayan Langurs ...
In response to dog predation, adult males performed three types of anti-predator response behaviors: direct fighting with a predator, emitting alarm calls, ...Missing: disturbances | Show results with:disturbances
[39]
(PDF) Social Organization and Reproductive Biology of Himalayan ...
Aug 6, 2025 · High resting and foraging in langurs even in the wild, might be attributed to a habitat condition with enough resources (Minhas et al., 2010; ...Missing: nesting | Show results with:nesting
[40]
Dominance relationships in male Nepal gray langurs ...
Feb 19, 2016 · Semnopithecus spp. vary in male group composition across populations, ranging from 100% one-male groups to 100% multimale groups (Koenig and ...
[41]
[PDF] PRIMATE REPORT PRIMATE REPORT - Deutsches Primatenzentrum
Two basic types of social groups are bisexual troops and all-male bands. Troops are matrilineal groups of adult females and offspring with either one adult ...<|separator|>
[42]
Infanticide as Sexual Conflict: Coevolution of Male Strategies and ...
Recent observations on resident male change followed by infanticide in Hanuman langurs (Semnopithecus entellus) around Jodhpur. ... Variation in langur social ...
[43]
Infanticide in hanuman langurs: social organization, male migration ...
Hanuman langurs (Presbytis entellus) have been among the first primate species where infanticide by adult males was observed and reported in the wild.
[44]
(PDF) Resource distribution and group size in the common ...
Aug 6, 2025 · They live in mixed-sex groups ranging from 2 to 27 individuals with a mean group size of 9.10 ± 5.54SD and a few all-male bands. The groups ...
[45]
[PDF] Genetic relatedness and socioecology of wild Hanuman langur ...
Across its range, it exhibits a variety of social organisations, from one-male multi female troops and all male bands, to multi-male, multi-female troops.Missing: scholarly | Show results with:scholarly
[46]
Paternity analysis of alternative male reproductive routes among the ...
In one-male groups all infants were sired by the only resident male in the group. In multimale groups the alpha male fathered significantly more infants (57%) ...Missing: care | Show results with:care
[47]
[PDF] Social Organization and Reproductive Biology of Himalayan Grey ...
Diurnal activities. The daily activities started from early morning and were always initiated by the resident male with morning “whoops”. After descending.
[48]
Hanuman Langur Presbytis (Semnopithecus) entellus
Gestation is from 190-210 days (Roonwal & Mohnot, 1977). Singleton births are the rule, but twins have also been observed. The weight at birth is around 500 g, ...Missing: period | Show results with:period
[49]
Tarai Gray Langur, Semnopithecus hector
They are overall a grayish-yellow color, with brown limbs, a gray-brown back, and a light gray belly, though overall, compared to other langur species, their ...Missing: features | Show results with:features
[50]
Evidence of Allomaternal Nursing across One-Male Units in ... - NIH
Jan 11, 2012 · Our findings suggest that lactating primate females are primed to care for young infants and, that the misdirected parental care hypothesis may ...
[51]
Hanuman langur (Semnopithecus entellus) longevity, ageing, and ...
Taxonomy: Kingdom: Animalia Phylum: Chordata Class: Mammalia (Taxon entry) Order: Primates (Taxon entry) Family: Cercopithecidae Genus: Semnopithecus; Species ...Missing: Groves | Show results with:Groves
[52]
VOCALIZATION IN HANUMAN LANGUR, SEMNOPITHECUS ...
Mar 3, 2004 · ... calls. Of these most common vocal repertoire is the loud calls or whoops, which are associated with preparation for movements, finalization ...
[53]
Predator mobbing by an all-male band of hanuman langurs ...
An all-male band of 15 Hanuman langurs alarm barked and mobbed a leopard in Sariska Tiger Reserve, a forest in Rajasthan, India. After initially alarm call.
[54]
[PDF] Eco-ethology of the common langur Semnopithecus entellus ...
Eco-ethology of the common langur Semnopithecus entellus (Dufresne) in ... Of the major diurnal activities viz., rest, move, sit, social groom, self ...
[55]
[PDF] social interactions of hanuman langur
Dec 24, 2014 · Dominant males are engaged in allogrooming to ensure protection during violent encounters, reconciliation of conflict within groups and ...
[56]
[PDF] The Communicative Functions of Touch in Humans, Nonhuman ...
Common touches in reconciliation include grooming, mounting, and clasping, which is ... Tactile communication among Free-ranging Langurs. American Journal of ...
[57]
[PDF] The evolution of female copulation calls in primates
Oct 11, 2005 · We hypothesize that female copulation calls have evolved under the selective pressures of risk of infanticide and sperm competition. When male ...
[58]
(PDF) Semnopithecus entellus - ResearchGate
PDF | On Sep 27, 2020, Honnavalli N. Kumara and others published Semnopithecus entellus | Find, read and cite all the research you need on ResearchGate.
[59]
Genetic Studies of Nilgiri Langur (<i>Semnopithecus johnii</i>) from ...
Sep 16, 2025 · The IUCN Red List has classified the Nilgiri langur as Vulnerable due to habitat loss and fragmentation of its forest habitat, and it continues ...
[60]
Habitat loss and poaching threaten the survival of the elusive Nilgiri ...
Jun 24, 2019 · The key threat faced by this langur, apart from habitat loss, is poaching. Nilgiri langurs are poached for their pelt, organs, blood and flesh.Missing: change | Show results with:change<|control11|><|separator|>
[61]
Surgical management of dental trauma and incisor teeth avulsion ...
Dec 21, 2022 · Surgical management of dental trauma and incisor teeth avulsion due to dog bite in a grey langur (Semnopithecus entellus) ... rabies among ...
[62]
Surveillance Opportunities and the Need for Intersectoral ... - Gale
There is a possibility of spill-over of rabies virus from dogs to wildlife and vice versa. ... langurs 48 Palm cat 41 Civet cat 14 Bandicoot 11 Bat 9 Wild ...
[63]
Studies gain pace while the identity of Kashmir's langurs remains gray
Dec 13, 2024 · “Climate change is likely to impact temperate deciduous forests, which are the primary habitat for the Kashmir gray langur,” Bashir explains. As ...Missing: poaching | Show results with:poaching
[64]
West to east shift in range predicted for Himalayan Langur in climate ...
The group of langur species in the Himalayan range, comprising Semnopithecus ajax, Semnopithecus hector and Semnopithecus schistaceus, now proved to be one ...
[65]
Appendices | CITES
The total weights present of each individual annotated species are considered individually to determine whether a CITES permit or certificate is required for ...
[66]
(PDF) Semnopithecus johnii, Nilgiri Langur THE IUCN RED LIST OF ...
Jul 23, 2020 · Semnopithecus johnii, Nilgiri Langur THE IUCN RED LIST OF THREATENED SPECIES™ ; Mewa Singh at University of Mysore · Mewa Singh · University of ...
[67]
A New Color Morph of the Southern Purple-faced Langur ... - BioOne
Feb 1, 2013 · The southern subspecies is primarily distinguishable by a defined silver-white rump patch; a detailed description of the standard pelage ...Missing: distinguishing | Show results with:distinguishing