Semnopithecus

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Semnopithecus
Gray langur (Semnopithecus) by Shantanu Kuveskar.jpg
Gray langur in Mangaon, Maharashtra
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Primates
Suborder: Haplorhini
Infraorder: Simiiformes
Family: Cercopithecidae
Subfamily: Colobinae
Tribe: Presbytini
Genus: Semnopithecus
Desmarest, 1822
Type species
Simia entellus [1]
Dufresne, 1797
Species

S. schistaceus
S. ajax
S. hector
S. entellus
S. hypoleucos
S. johnii
S. priam
S. vetulus

Contents

Semnopithecus is a genus of Old World monkeys native to the Indian subcontinent, with all species with the exception of two being commonly known as gray langurs. [1] Traditionally only one species Semnopithecus entellus was recognized, but since about 2001, additional species have been recognized. The taxonomy has been in flux, but currently eight species are recognized.

Members of the genus Semnopithecus are terrestrial, inhabiting forest, open lightly wooded habitats, and urban areas on the Indian subcontinent. Most species are found at low to moderate altitudes, but the Nepal gray langur and Kashmir gray langur occur up to 4,000 m (13,000 ft) in the Himalayas. [2] [3]

Characteristics

These langurs are largely gray (some more yellowish), with a black face and ears. Externally, the various species mainly differ in the darkness of the hands and feet, the overall color and the presence or absence of a crest. [4] [5] Typically all north Indian gray langurs have their tail tips looping towards their head during a casual walk whereas all south Indian and Sri Lankan gray langurs have an inverted "U" shape or a "S" tail carriage pattern. [6] There are also significant variations in the size depending on the sex, with the male always larger than the female. The head-and-body length is from 51 to 79 cm (20 to 31 in). Their tails, at 69 to 102 cm (27 to 40 in) are always longer than their bodies. [7] Langurs from the southern part of their range are smaller than those from the north. At 26.5 kg (58 lb), the heaviest langur ever recorded was a male Nepal gray langur. [5] The larger gray langurs are rivals for the largest species of monkey found in Asia. The average weight of gray langurs is 18 kg (40 lb) in the males and 11 kg (24 lb) in the females. [7]

Langurs mostly walk quadrupedally and spend half of their time on the ground and the other half in trees. They will also make bipedal hops, climbing and descending supports with the body upright, and leaps. Langurs can leap 3.6–4.7 m (12–15 ft) horizontally and 10.7–12.2 m (35–40 ft) in descending. [8]

Taxonomy

Young gray langur at Nagarhole National Park, Mysore Gray Langur - Young.jpg
Young gray langur at Nagarhole National Park, Mysore
Hanuman langur in Nandankanan Zoological Park in Bhubaneshwar Hanuman Langur or Gray Langur.jpg
Hanuman langur in Nandankanan Zoological Park in Bhubaneshwar

Traditionally, only Semnopithecus entellus was recognized as a species, the remainder all being treated as subspecies. In 2001, it was proposed that seven species should be recognized. [4] This was followed in Mammal Species of the World in 2005, [1] though several of the seven species intergrade, and alternative treatments exist where only two species (a northern and a southern) are recognized. [5] Phylogenetic evidence supports at least three species: a north Indian, a south Indian and a Sri Lankan one. [9] [10]

It has been suggested that the Semnopithecus priam thersites is worthy of treatment as a species rather than a subspecies, but at present this is based on limited evidence. [11] During a study based on external morphology and ecological niche modelling in Peninsular India six main types were found, but continued to label all as subspecies. Coat color is highly variable, possible due to phenotypic plasticity and therefore of questionable value in species delimitation. [12] [13]

It has been suggested that Trachypithecus should be considered only a subgenus of Semnopithecus. [5] If maintaining the two as separate monophyletic genera, the purple-faced langur and Nilgiri langur belong in Semnopithecus instead of their former genus Trachypithecus. At present it is unclear where the T. pileatus species group (consisting of the capped langur, Shortridge's langur and Gee's golden langur) belongs, as available mtDNA data place it in Semnopithecus, while Y chromosome data place it in Trachypithecus. [9] A possible explanation for this is that the T. pileatus species group is the result of fairly recent hybridization between Semnopithecus and Trachypithecus. [10]

As of 2005, the authors of Mammal Species of the World recognized the following seven Semnopithecus species [1]

Results of analysis of mitochondrial cytochrome b gene and two nuclear DNA-encoded genes of several colobine species revealed that Nilgiri and purple-faced langurs cluster with gray langur, while Trachypithecus species form a distinct clade. [14] Since then, two other species have been moved from Trachypithecus to Semnopithecus: [15] [16]

In addition, Semnopithecus dussumieri has been determined to be invalid. [12] [16] [17] [18] Most of the range that had been considered S. dussumieri is now considered S. entellus. [15] [16]

Thus the current generally accepted species within the genus Semnopithecus are: [16] [17] [18]

Genus Semnopithecus Desmarest, 1822 – eight species
Common nameScientific name and subspeciesRangeSize and ecologyIUCN status and estimated population
Black-footed gray langur

Black Footed Langur Log Nagarhole Dec21 D72 21387.jpg

S. hypoleucos
Blyth, 1841

Three subspecies
  • S. h. achates
  • S. h. hypoleucos
  • S. h. iulus
Southern India
Black-footed Gray Langur area.png
Size: 41–78 cm (16–31 in) long, plus 69–108 cm (27–43 in) tail [19]

Habitat: Forest and shrubland [20]

Diet: Leaves, fruit, and flowers [19]
 LC 


Unknown Decrease2.svg [20]

Kashmir gray langur


S. ajax
Pocock, 1928
Himalayas
Kashmir Gray Langur area.png
Size: 41–78 cm (16–31 in) long, plus 69–108 cm (27–43 in) tail [19]

Habitat: Forest [3]

Diet: Leaves, bark, and seeds [3]
 EN 


1,400–1,500 Decrease2.svg [3]

Nepal gray langur

Nepal gray langur, Bhutan.jpg

S. schistaceus
Hodgson, 1840
Himalayas
Nepal Gray Langur area.png
Size: 41–78 cm (16–31 in) long, plus 69–108 cm (27–43 in) tail [19]

Habitat: Forest, shrubland, and rocky areas [2]

Diet: Leaves and fruit, as well as seeds, roots, flowers, bark, twigs, coniferous cones, moss, lichens, ferns, shoots, rhizomes, grass, and invertebrate animals [2]
 LC 


Unknown Decrease2.svg [2]

Nilgiri langur

Nilgiri langur (1) by N. A. Naseer.jpg

S. johnii
(J. Fischer, 1829)

Nilgiri Langur area.png
Size: 41–78 cm (16–31 in) long, plus 69–108 cm (27–43 in) tail [19]

Habitat: Forest [21]

Diet: Leaves, fruit, and flowers [19]
 VU 


9,500–10,000 Steady2.svg [21]

Northern plains gray langur

Northern plains gray langur.jpg

S. entellus
(Dufresne, 1797)
India
Bengal Gray Langur area.png
Size: 41–78 cm (16–31 in) long, plus 69–108 cm (27–43 in) tail [19]

Habitat: Forest, savanna, and shrubland [22]

Diet: Leaves, fruit, and flowers, as well as insects, bark, gum, and soil [23]
 LC 


Unknown Decrease2.svg [22]

Tarai gray langur

Tarai Grey Langur.jpg

S. hector
Pocock, 1928
Himalayas
Tarai Gray Langur area.png
Size: 41–78 cm (16–31 in) long, plus 69–108 cm (27–43 in) tail [19]

Habitat: Forest [24]

Diet: Leaves, fruit, and flowers [19]
 NT 


Unknown Decrease2.svg [24]

Tufted gray langur

Tufted grey langur (Semnopithecus priam).jpg

S. priam
Blyth, 1844

Three subspecies
  • S. p. anchises
  • S. p. priam
  • S. p. thersites
Southern India and Sri Lanka
Tufted Gray Langur area.png
Size: 41–78 cm (16–31 in) long, plus 69–108 cm (27–43 in) tail [19]

Habitat: Forest and shrubland [25]

Diet: Leaves and fruit [25]
 NT 


Unknown Decrease2.svg [25]

Purple-faced langur

Purple-faced langur (Semnopithecus vetulus vetulus).jpg

S. vetulus
(Erxleben, 1777)

Four subspecies
  • T. v. monticola (Montane purple-faced langur)
  • T. v. nestor (Western purple-faced langur)
  • T. v. philbricki (Dryzone purple-faced langur)
  • T. v. vetulus (Southern lowland wetzone purple-faced langur)
Sri Lanka
Purple-faced Langur area.png
Size: 41–78 cm (16–31 in) long, plus 69–108 cm (27–43 in) tail [19]

Habitat: Forest [26]

Diet: Leaves, fruit, flowers, and seeds [26]
 EN 


Unknown Decrease2.svg [26]

A 2013 genetic study indicated that while S. entellus, S. hypoleucos, S. priam and S. johnii are all valid taxa, there has been hybridization between S. priam and S. johnii. [27] It also indicated that there has been some hybridization between S. entellus and S. hypoleucos where their ranges overlap, and a small amount of hybridization between S. hypoleucos and S. priam. [27] It also suggested that S. priam and S. johnii diverged from each other fairly recently. [27]

Distribution and habitat

The entire distribution of all gray langur species stretches from the Himalayas in the north to Sri Lanka in the south, and from Bangladesh in the east to Pakistan in the west. [28] They possibly occur in Afghanistan. [5] The bulk of the gray langur distribution is within India, and all seven currently recognized species have at least a part of their range in this country. [4]

Gray langurs can adapt to a variety of habitats. [29] They inhabit arid habitats like deserts, tropical habitats like tropical rainforests and temperate habitats like coniferous forests, deciduous habitats and mountains habitats. They are found at sea level to altitudes up to 4,000 m (13,000 ft). [2] [3] They can adapt well to human settlements, and are found in villages, towns and areas with housing or agriculture. [30] They live in densely populated cities like Jodhpur, which has a population numbering up to a million. [31]

Ecology and behavior

Gray langur GREY LANGUR.jpg
Gray langur

Gray langurs are diurnal. They sleep during the night in trees but also on man-made structures like towers and electric poles when in human settlements. [32] When resting in trees, they generally prefer the highest branches. [33]

Ungulates like bovine and deer will eat food dropped by foraging langurs. [34] Langurs are preyed upon by leopards, dholes and tigers. [35] Wolves, jackals, Asian black bears and pythons may also prey on langurs.

Diet

Gray Langur near a village in Rajasthan Gary Langur, RJ IN 2011.jpg
Gray Langur near a village in Rajasthan

Gray langurs are primarily herbivores. However, unlike some other colobines they do not depend on leaves and leaf buds of herbs, but will also eat coniferous needles and cones, fruits and fruit buds, evergreen petioles, shoots and roots, seeds, grass, bamboo, fern rhizomes, mosses, and lichens. Leaves of trees and shrubs rank at the top of preferred food, followed by herbs and grasses. Non-plant material consumed include spider webs, termite mounds and insect larvae. [36] They forage on agricultural crops and other human foods, and even accept handouts. [37] Although they occasionally drink, langurs get most of their water from the moisture in their food. [38]

Social structure

Juvenile gray langur Juvenile Indian Grey Langur.jpg
Juvenile gray langur

Gray langurs exist in three types of groups:

All-male groups tend to be the smallest of the groups and can consist of adults, subadults, and juveniles. Some populations have only multiple-male groups as mixed sex groups, while others have only one-male groups as mixed sexed groups. [39]

Some evidence suggests multiple-male groups are temporary and exist only after a takeover, and subsequently split into one-male and all-male groups. [41]

Social hierarchies exist for all group types. [42] [43] In all-male groups, dominance is attained through aggression and mating success. [44] With sexually mature females, rank is based on physical condition and age. [45] [46] The younger the female, the higher the rank. Dominance rituals are most common among high-ranking langurs. [43] Most changes in social rank in males take place during changes in group members. An adult male may remain in a one-male group for 45 months. [47] The rate of male replacement can occur quickly or slowly depending on the group. [48]

Females within a group are matrilineally related. Female memberships are also stable, but less so in larger groups. [40] [45] Relationships between the females tend to be friendly. They will do various activities with each together, such as foraging, traveling and resting. They will also groom each other regardless of their rank. However, higher-ranking females give out and receive grooming the most. [49] In addition, females groom males more often than the other way around. [50] Male and female relationships are usually positive. Relationships between males can range from peaceful to violent. While females remain in their natal groups, males will leave when they reach adulthood. [40] Relationships between groups tend to be hostile. High-ranking males from different groups will display, vocalize, and fight among themselves.

Reproduction and parenting

Tufted gray langur with young Semnopithecus priam.jpg
Tufted gray langur with young
Gray langur with newborn GrayLangurWithNewBorn1.JPG
Gray langur with newborn

In one-male groups, the resident male is usually the sole breeder of the females and sires all the young. In multiple-male groups, the highest-ranking male fathers most of the offspring, followed by the next-ranking males and even outside males will father young. [51] Higher-ranking females are more reproductively successful than lower-ranking ones. [46]

Female gray langurs do not make it obvious that they are in estrous. However, males are still somehow able to deduce the reproduction state of females. [52] Females signal that they are ready to mate by shuddering the head, lowering the tail, and presenting their anogenital regions. [53] Such solicitations do not always lead to copulation. When langurs mate, they are sometimes disrupted by other group members. [47] Females have even been recorded mounting other females. [54]

The gestation period of gray langur lasts around 200 days, at least at Jodhpur, India. In some areas, reproduction is year-around. [53] Year-round reproduction appears to occur in populations that capitalize on human-made foods. Other populations have seasonal reproduction. [47] Infanticide is common among gray langurs. Most infanticidal langurs are males that have recently immigrated to a group and driven out the prior male. These males only kill infants that are not their own. [55] Infanticide is more commonly reported in one-male groups, perhaps because one male monopolizing matings drives the evolution of this trait. In multiple-male groups, the costs for infanticidal males are likely to be high as the other males may protect the infants and they can't ensure that they'll sire young with other males around. Nevertheless, infanticide does occur in these groups, and is suggested that such practices serve to return a female to estrous and gain the opportunity to mate. [56]

Females usually give birth to a single infant, although twins do occur. Most births occur during the night. [57] Infants are born with thin, dark brown or black hair and pale skin. Infants spend their first week attached to their mothers' chests and mostly just suckle or sleep. [58] They do not move much in terms of locomotion for the first two weeks of their life. As they approach their sixth week of life, infants vocalize more. [59] They use squeaks and shrieks to communicate stress. In the following months, the infants are capable of quadrupedal locomotion and can walk, run and jump by the second and third months. Alloparenting occurs among langurs, starting when the infants reach two years of age. The infant will be given to the other females of the group. However, if the mother dies, the infant usually follows. [58] Langurs are weaned by 13 months.

Vocalizations

Gray langurs are recorded to make a number of vocalizations: [60] [61]

Status and conservation

Gray langurs have stable populations in some areas and declining ones in others. [62] Both the black-footed gray langur and Kashmir gray langur are considered threatened. [3] [20] The latter is the rarest species of gray langur, with less than 250 mature individuals remaining. [3]

In India, gray langurs number at around 300,000. [63] India has laws prohibiting the capturing or killing of langurs, but they are still hunted in some parts of the country. [12] Enforcement of these laws has proven to be difficult and it seems most people are unaware of their protection. [64] Populations are also threatened by mining, forest fires and deforestation for wood. [65]

Langurs can be found near roads and can become victims of automobile accidents. This happens even in protected areas, with deaths by automobile collisions making nearly a quarter of mortality in Kumbhalgarh Wildlife Sanctuary in Rajasthan, India. [66] Langurs are considered sacred in the Hindu religion and are sometimes kept for religious purposes by Hindu priests and for roadside performances. However, some religious groups use langurs as food and medicine, and parts of gray langurs are sometimes kept as amulets for good luck. [67]

Because of their sacred status and their less aggressive behavior compared to other primates, langurs are generally not considered pests in many parts of India. Despite this, research in some areas show high levels of support for the removal of langurs from villages, their sacred status no longer important. [68] Langurs will raid crops and steal food from houses, and this causes people to persecute them. [68] While people may feed them in temples, they do not extend such care to monkeys at their homes. [69] Langurs stealing and biting people to get food in urban areas may also contribute to more persecutions. [70]

See also

Related Research Articles

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The purple-faced langur, also known as the purple-faced leaf monkey, is a species of Old World monkey that is endemic to Sri Lanka. The animal is a long-tailed arboreal species, identified by a mostly brown appearance, dark face and a very shy nature. The species was once highly prevalent, found in suburban Colombo and the "wet zone" villages, but rapid urbanization has led to a significant decrease in the population level of the monkeys. It had traditionally been classified within the genus Trachypithecus but was moved to the genus Semnopithecus based on DNA evidence indicating that is it more closely related to the gray langurs.

<span class="mw-page-title-main">Colobinae</span> Subfamily of Old World monkeys

The Colobinae or leaf-eating monkeys are a subfamily of the Old World monkey family that includes 61 species in 11 genera, including the black-and-white colobus, the large-nosed proboscis monkey, and the gray langurs. Some classifications split the colobine monkeys into two tribes, while others split them into three groups. Both classifications put the three African genera Colobus, Piliocolobus, and Procolobus in one group; these genera are distinct in that they have stub thumbs. The various Asian genera are placed into another one or two groups. Analysis of mtDNA confirms the Asian species form two distinct groups, one of langurs and the other of the "odd-nosed" species, but are inconsistent as to the relationships of the gray langurs; some studies suggest that the gray langurs are not closely related to either of these groups, while others place them firmly within the langur group.

<i>Trachypithecus</i> Genus of Old World monkeys

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<i>Presbytis</i> Genus of south-east Asian monkeys

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<span class="mw-page-title-main">Gee's golden langur</span> Species of Old World monkey

Gee's golden langur, also known as simply the golden langur, is an Old World monkey found in a small region of Western Assam, India and in the neighboring foothills of the Black Mountains of Bhutan. Long considered sacred by many Himalayan people, the golden langur was first brought to the attention of the Western world by the naturalist Edward Pritchard Gee in the 1950s. Adult males have a cream to golden coat with darker flanks while the females and juveniles are lighter. The golden langur has a black face and a long tail up to 50 cm (19.69 in) in length. It lives in high trees and has a herbivorous diet of fruits, leaves, seeds, buds and flowers. The average group size is eight individuals, with a ratio of several females to each adult male. It is one of the most endangered primate species of India and Bhutan.

<span class="mw-page-title-main">Nilgiri langur</span> Species of Old World monkey

The Nilgiri langur is a langur. This primate has glossy black fur on its body and golden brown fur on its head. It is similar in size and long-tailed like the gray langurs. Females have a white patch of fur on the inner thigh. It typically lives in troops of nine to ten monkeys. Its diet consists of fruits, shoots and leaves. The species is classified as vulnerable due to habitat destruction and poaching for its fur and flesh, the latter believed to have aphrodisiac properties.

<span class="mw-page-title-main">Black-footed gray langur</span> Species of Old World monkey

The black-footed gray langur is an Old World monkey, one of the species of langurs. Found in southern India, this, like other gray langurs, is a leaf-eating monkey.

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<span class="mw-page-title-main">Northern plains gray langur</span> Species of mammal

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<span class="mw-page-title-main">Dusky leaf monkey</span> Species of Old World monkey

The dusky leaf monkey, also known as the spectacled langur or the spectacled leaf monkey, is a species of primate in the family Cercopithecidae. It is found in Peninsular Malaysia, Myanmar and Thailand, and can occasionally be found in Singapore. During the day, these small, folivorous primates divide in sub-groups and forage for vegetation and fruit throughout the tropical forests. According to the IUCN, the dusky leaf monkey's population is declining due to habitat loss, poaching, and anthropogenic land use, which prompted the IUCN to classify the species as endangered in 2015.

<span class="mw-page-title-main">Southern plains gray langur</span>

Southern plains gray langur was the common name ascribed to Semnopithecus dussumieri by Mammal Species of the World in 2005. Along with several other Semnopithecus, it had been previously considered a subspecies of the northern plains gray langur, Semnopithecus entellus, i.e., Semnopithecus entellus dussumieri. Subsequent genetic research has revealed that Semnopithecus dussumieri is an invalid taxon. These monkeys live in groups in forests and other rural habitats, with some groups being habituated to human contact and feeding. They are herbivorous, feeding by day mainly on foliage, fruits and flowers, and sleeping at night high in a tree.

<span class="mw-page-title-main">Nepal gray langur</span> Species of Old World monkey

The Nepal gray langur is a gray langur endemic to the Himalayas in Nepal, far southwestern Tibet, northern India, northern Pakistan, Bhutan and possibly Afghanistan. It is found in forests at an elevation of 1,500 to 4,000 metres. Its easternmost limit in India is Buxa Tiger Reserve in northern West Bengal, at least up to the Rydak river.

<span class="mw-page-title-main">Kashmir gray langur</span> Species of Old World monkey

The Kashmir gray langur is an Old World monkey, one of the langur species. It is a leaf-eating monkey.

<span class="mw-page-title-main">Tufted gray langur</span> Species of Old World monkey

The tufted gray langur, also known as Madras gray langur, and Coromandel sacred langur, is an Old World monkey, one of the species of langurs. This, like other gray langurs, is mainly a leaf-eating monkey. It is found in southeast India and Sri Lanka. It is one of three Semnopithecus species named after characters from The Iliad, S. hector and S. ajax being the others. In Sinhala it is known as හැලි වදුරා.

<span class="mw-page-title-main">Western purple-faced langur</span> Subspecies of Old World monkey

The western purple-faced langur, also known as the north lowland wetzone purple-faced langur, is a subspecies of purple-faced langur endemic to Sri Lanka. It lives in the wet zone in western Sri Lanka around the former capital city of Colombo. This subspecies are known to live in tropical rainforest, specifically in an area of Sri Lanka, where it is noted to rain two times more compared to the neighboring region. The subspecies is generally gray-brown with lighter whiskers, a gray rump patch, and dark forearms and legs. These monkeys also have dark brown or black, hairless faces. Their hands and feet are also hairless. When infants, their fur is medium brown to light gray, however it changes to their darker adult colors at between 12 and 16 weeks old. Typical length is between 48 and 67 centimetres excluding tail, with a 59 to 85 centimetres tail. On average, males weigh 8.5 kilograms (19 lb) and females weigh 7.8 kilograms (17 lb). Their average life span is about 26 years if the monkey was held in captivity. Due to their leafy diet, the western purple-faced langur back teeth, include high crowns and pointed cusps to make eating through greenery simpler. They also developed a digestive system that would allow them to break down cellulose in the leaves they eat. Although, leaves are their primary source of food, they also eat fruits and flowers. Female langurs hold a pregnancy for 195–210 days, and after the birth the offsprings are taken closely cared for by the mother until they are about 12–20 weeks old.

Infanticide in non-human primates occurs when an individual kills its own or another individual's dependent young. Five hypotheses have been proposed to explain infanticide in non-human primates: exploitation, resource competition, parental manipulation, sexual selection, and social pathology.

<span class="mw-page-title-main">Selangor silvered langur</span> Species of Old World monkey

The Selangor silvered langur is a species of leaf monkey found on the west coast of Peninsular Malaysia. It had been previously considered a form of silvery lutung. Roos and colleagues elevated this population to a subspecies level, Trachypithecus cristatus selangorensis, in 2008. It has since come to be regarded by primatologists as a separate species, Trachypithecus selangorensis.

<span class="mw-page-title-main">Presbytini</span> Tribe of Old World monkeys

Presbytini is a tribe of Old World monkeys that includes all of the Asian colobine monkeys.

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