Trachypithecus is a genus of Old World monkeys containing species known as lutungs, langurs, or leaf monkeys. Their range is much of Southeast Asia (northeast India, Vietnam, southern China, Borneo, Thailand, Java, and Bali).

The name "lutung" comes from the Sundanese language meaning "blackness", ultimately from Proto-Austronesian *luCuŋ (which originally referred to the Formosan rock macaque); it is preferred in one paper because the authors wanted the name langurs to only refer to monkeys in the genus Semnopithecus, although some "lutungs" are now "langurs" again. The scientific name of the genus comes from the Ancient Greek τραχύς (trakhús), meaning "rough", and πίθηκος (píthēkos), meaning "monkey".

Evolution

Trachypithecus cristatus robustus skull

Genetic analysis indicates that the ancestors of the modern species of lutung first differentiated from one another a little over 3 million years ago, during the late Pliocene. The various species alive today then diverged during the Pleistocene, presumably driven by habitat changes during the Ice Ages. The oldest fossils clearly identified as belonging to the genus date from the middle Pleistocene of Vietnam and Laos; later fossils are also known from Thailand, Java, and Sumatra. The closest living relatives of the lutungs are probably either the gray langurs or the surilis, although the exact relationships remain unclear, possibly due to hybridisation between these genera during the course of their recent evolutionary history.

Taxonomy

Nilgiri langur, formerly classified within the genus Trachypithecus but since moved to the genus Semnopithecus

As of 2005, the authors of Mammal Species of the World recognize the following Trachypithecus species:

Since then, the T. vetulus group (the purple-faced langur and the Nilgiri langur) have been moved the genus Semnopithecus based on DNA and other evidence.

In 2008, Roos et al. described the Malay Peninsula form of the silvery lutung as a separate subspecies and subsequently it has been elevated to a separate species within the T. cristatus group as the Selangor silvered langur, T. selangorensis. Roos et al. also elevated the West Javan Langur, Trachypithecus mauritius, and Annamese Langur, Trachypithecus margarita, to species status (formerly subspecies of T. auratus and T. germaini, respectively). In 2020, Roos et al. discovered a new species, Popa langur (T. popa), which is found only in Myanmar. Lastly, the White-headed langur (T. leucocephalus), previously thought to be a subspecies of the Francois langur (T. Francois) or Cat Ba langur (T. poliocephalus), is currently recognized as a distinct species by IUCN Red List assessors and the American Society of Mammalogists, based on a 2007 paper by Groves.

This leaves the current understanding of the genus Trachypithecus to be:

Genus Trachypithecus – Reichenbach, 1862 – twenty-one species
Common nameScientific name and subspeciesRangeSize and ecologyIUCN status and estimated population
Annamese langurT. margarita (Elliot, 1909)Southeastern AsiaSize: 40–76 cm (16–30 in) long, plus 57–110 cm (22–43 in) tail Habitat: Forest Diet: Leaves, flowers, and fruitEN Unknown
Capped langurT. pileatus (Blyth, 1843) Three subspecies T. p. brahmaT. p. pileatusT. p. tenebricusSouthern AsiaSize: 40–76 cm (16–30 in) long, plus 57–110 cm (22–43 in) tail Habitat: Forest Diet: Leaves, seeds, and fruit, as well as flowers, buds, bark, and caterpillarsVU Unknown
Cat Ba langurT. poliocephalus (Pousargues, 1898)Cát Bà Island, Vietnam (in purple)Size: 40–76 cm (16–30 in) long, plus 57–110 cm (22–43 in) tail Habitat: Forests and caves Diet: Leaves, flowers, and fruitCR 30–35
Delacour's langurT. delacouri (Osgood, 1911)Northern VietnamSize: 40–76 cm (16–30 in) long, plus 57–110 cm (22–43 in) tail Habitat: Forest, rocky areas, and caves Diet: Leaves, flowers, and fruitCR 240–250
Dusky leaf monkeyT. obscurus (Reid, 1837) Seven subspecies T. o. carboT. o. flavicaudaT. o. haloniferT. o. obscurusT. o. sanctorumT. o. seimundiT. o. styxSoutheastern AsiaSize: 42–61 cm (17–24 in) long, plus 50–85 cm (20–33 in) tail Habitat: Forest Diet: Leaves, shoots, fruit, and seedlingsEN Unknown
East Javan langurT. auratus (É. Geoffroy, 1812)Java and nearby islands in IndonesiaSize: 44–65 cm (17–26 in) long, plus 61–87 cm (24–34 in) tail Habitat: Forest Diet: Leaves and flowers, as well as fruit and insect larvaeVU Unknown
François' langurT. francoisi (Pousargues, 1898)Southern AsiaSize: 40–76 cm (16–30 in) long, plus 57–110 cm (22–43 in) tail Habitat: Forest, rocky areas, and caves Diet: Leaves, fruit, and seeds, as well as insectsEN 2,000–2,100
Gee's golden langurT. geei (Khajuria, 1956)Southern AsiaSize: 50–75 cm (20–30 in) long, plus 70–100 cm (28–39 in) tail Habitat: Forest Diet: Fruit, leaves, flowers, seeds, and twigsEN 6,000–6,500
Germain's langurT. germaini (A. Milne-Edwards, 1876)Southeastern AsiaSize: 40–76 cm (16–30 in) long, plus 57–110 cm (22–43 in) tail Habitat: Forest and rocky areas Diet: Leaves, fruit, and flowersEN Unknown
Hatinh langurT. hatinhensis (Dao, 1970)VietnamSize: 40–76 cm (16–30 in) long, plus 57–110 cm (22–43 in) tail Habitat: Forest, rocky areas, and caves Diet: Leaves, as well as fruit, vines, and flowersEN Unknown
Indochinese black langurT. ebenus (Brandon-Jones, 1995)Southeastern AsiaSize: 40–76 cm (16–30 in) long, plus 57–110 cm (22–43 in) tail Habitat: Forest, rocky areas, and caves Diet: Leaves, as well as fruit, vines, and flowersEN Unknown
Indochinese grey langurT. crepuscula (Elliot, 1909)Southeast Asia (in red)Size: 40–76 cm (16–30 in) long, plus 57–110 cm (22–43 in) tail Habitat: Forest, inland wetlands, and rocky areas Diet: Leaves, flowers, and fruitEN 2,400–2,500
Laotian langurT. laotum (Thomas, 1911)LaosSize: 40–76 cm (16–30 in) long, plus 57–110 cm (22–43 in) tail Habitat: Forest and rocky areas Diet: Leaves, flowers, and fruitEN Unknown
Phayre's leaf monkeyT. phayrei (Blyth, 1847) Two subspecies T. p. phayreiT. p. shanicusSoutheast Asia (in green)Size: 40–76 cm (16–30 in) long, plus 57–110 cm (22–43 in) tail Habitat: Forest Diet: Leaves, as well as bamboo shootsEN Unknown
Popa langurT. popa Roos et al., 2020MyanmarSize: 40–76 cm (16–30 in) long, plus 57–110 cm (22–43 in) tail Habitat: Forest Diet: Leaves, flowers, and fruitCR 130–180
Shortridge's langurT. shortridgei (Wroughton, 1915)Southern AsiaSize: 40–76 cm (16–30 in) long, plus 57–110 cm (22–43 in) tail Habitat: Forest Diet: Leaves, flowers, and fruitEN Unknown
Selangor silvered langurT. selangorensis Roos, Nadler, Walter, 2008Peninsular MalaysiaSize: 40–76 cm (16–30 in) long, plus 57–110 cm (22–43 in) tail Habitat: Forest Diet: Leaves, flowers, and fruitNT Unknown
Silvery lutungT. cristatus (Raffles, 1821) Two subspecies T. c. cristatusT. c. vigilansSoutheastern AsiaSize: 46–56 cm (18–22 in) long, plus 63–84 cm (25–33 in) tail Habitat: Forest Diet: Leaves, as well as fruit, seeds, shoots, flowers, and budsVU Unknown
Tenasserim lutungT. barbei (Blyth, 1847)Southeastern AsiaSize: 40–76 cm (16–30 in) long, plus 57–110 cm (22–43 in) tail Habitat: Forest Diet: Leaves, flowers, and fruitVU Unknown
West Javan langurT. mauritius (Griffith, 1821)Island of JavaSize: 40–76 cm (16–30 in) long, plus 57–110 cm (22–43 in) tail Habitat: Forest Diet: Leaves, flowers, and fruitVU Unknown
White-headed langurT. leucocephalus Tan, 1957Southern ChinaSize: 40–76 cm (16–30 in) long, plus 57–110 cm (22–43 in) tail Habitat: Rocky areas Diet: Leaves, flowers, and fruitCR 230–250

Physical description

Francois' langur (Trachypithecus francoisi)

Lutungs have a rather slim build with a long tail. The fur color varies, depending on the species, from black and grey to orange yellow. Many species have skin designs and a brighter lower surface, the hair on the head is often compared to a hood. Their arms are very short in comparison to their hind legs, and their thumbs are also somewhat shorter than in other primates. The inner surfaces of the hands and feet are hairless so that their fur does not get caught when reaching into branches. These animals reach a length of 40 to 80 cm and a weight of 5 to 15 kg, with males being generally larger than females. A ridge over the eyes and other details, primarily related to the head, differentiate the lutungs from the surilis.

Habitat and distribution

Lutungs live in forests. They often prefer rainforests, although they are occasionally also found in secluded mountain forests and limestone karst forests. Lutungs are found in South-east Asia and parts of South Asia, from India in the west to China in the east.

Behaviour

Lutungs spend the largest part of the day in the trees, where they walk along the branches on all fours.They also jump from tree to tree, often covering long distances between trees; those long jumps are referred to as leaping. Lutungs are diurnal, although more active in the early mornings and the afternoon.

They live in groups of five to 20 animals, mostly in harems, i.e. a single male with several females. Young males must leave their birth group when fully mature, often forming bachelor groups. If a new male takes over a harem, defeating or scaring off the former harem leader, he often kills the infants in the group. This behavior is also known as infanticide. Lutungs are territorial, and emit loud calls to defend their territories from rival males, resorting to force if the outsiders are not scared off. They have a common repertoire of sounds with which they warn group members. Mutual grooming also plays an important role in maintaining the bonds between individuals.

Lutungs are herbivores, primarily eating leaves, fruits, and buds. To digest the tough leaves, they developed a multichambered stomach.

Reproduction

Lutungs typically give birth to a single young, after a seven-month gestation period. Twins occur, but are rare. Newborns usually have golden-yellow or orange fur. The mother shares responsibilities of rearing the young with the other females ("aunties") of the harem. Females within a group hand the young around, play with them, carry them, and cuddle them, allowing the mother to forage or spend time alone. If the mother dies, another female adopts the young animal. Lutungs are weaned in the latter half of their first year, and reach full maturity at 4 to 5 years. The life expectancy is estimated at 20 years.

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