Gray slender loris

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Gray slender loris
Slender Loris.jpg
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Primates
Suborder: Strepsirrhini
Family: Lorisidae
Genus: Loris
Species:
L. lydekkerianus [2]
Binomial name
Loris lydekkerianus [2]
Cabrera, 1908
Subspecies

L. l. lydekkerianus(Cabrera, 1908)
L. l. malabaricus(Wroughton, 1917)
L. l. nordicus(Osman Hill, 1933)
L. l. grandis(Osman Hill and Phillips, 1932)

Contents

Gray Slender Loris area.png
Gray slender loris range

The gray slender loris (Loris lydekkerianus) is a species of primate in the family Loridae. It is native to India and Sri Lanka and inhabits subtropical and tropical dry forests and subtropical or tropical moist lowland forests. It is threatened by habitat loss.

Taxonomy

Together with the red slender loris (Loris tardigradus), the grey slender loris (Loris lydekkerianus) is a type of slender loris (genus Loris) in the strepsirrhine primate family Lorisidae. [2] In 1908 Spanish zoologist Ángel Cabrera first described the Mysore slender loris (Loris tardigradus lydekkerianus) in Chennai, India, which he named for the English naturalist Richard Lydekker. This subspecies was further described by William Charles Osman Hill in his seminal primate book Primates: Comparative Anatomy and Taxonomy (1953). Hill believed there was one species of slender loris which was further split into six subspecies, two in India and four in Sri Lanka. [3] In 1998 biological anthropologist Colin Groves recognised two species, L. tardigradus and L. lydekkerianus, [4] which have been widely accepted by the scientific community. [5] [6] [7] Loris lydekkerianus now includes four geographically separated subspecies, L. l. lydekkerianus (previously L. t. lydekkerianus) and L. l. malabaricus in India and L. l. nordicus and L. l. grandis in Sri Lanka. [8]

A slender loris group known as the montane slender loris (taxon nycticeboides) has had uncertain classification and variously placed as a subspecies of L. lydekkerianus, L. tardigradus, and as a distinct species. [5] [9] A 2019 study based on partial CO1 sequences showed the taxon can be classified in a single haplogroup with L. t. tardigradus. [10]

Physical description

Skull Loris lydekkerianus skull.jpg
Skull

Slender lorises have extremely gracile limbs and extreme stereoscopic vision. The gray slender loris has a wide variation in pelage colour and each subspecies can be identified by this. The fur is short and gray or reddish on their backs, sometimes a darker stripe extends from the top of their head to the end of their back. The ventrum is white or buff-coloured. The eyes are surrounded with darker fur and have orbits which look straight forward giving excellent stereoscopic vision and are located the closest together amongst the primates. Between the eyes there is a white patch of hair known as the median stripe. The muzzle is larger and less pointed than the red slender loris. The hands are also highly specialised with a reduced second digit that allows the loris to cling to small branches. This is further augmented by adaptions in the first metatarsal that are thought to allow an extremely firm grasp. The tail is vestigial. The average head-body length is 18–26 centimetres (7.1–10.2 in). [11]

In Sri Lankan subspecies, Loris lydekkerianus grandis has short ears, and a heart-shaped face. Basal hairs of the vent of Loris lydekkerianus grandis are black and whereas those of Loris lydekkerianus nordicus are white in colour.

Behaviour

The gray slender loris is nocturnal and arboreal. Loris lydekkerianus nordicus 003.jpg
The gray slender loris is nocturnal and arboreal.

The behaviour of the gray slender loris is amongst the least known of the primates, despite the relatively large number of studies undertaking during the 2000s. [12] Like other lorises, they are nocturnal and emerge from their roost cavities only at dusk. They are mainly insectivorous. They primarily eat insects but do occasionally eat fruits, flowers, and small animals like mice and geckos when given the opportunity. [13] In southern India, the nominate race is often found in acacia and tamarind dominated forests or scrubs near cultivations. [14] Males hold larger home ranges than females. They are usually solitary while foraging, and it is rare for them to be seen in pairs or groups. However they may roost in groups of up to 7 [1] that include young of the recent and older litters. Adult males and females have individual home ranges and sleeping group associations are usually composed of a female and her offspring. [12] Communicate with a range of vocalisations and also use urine and scent marking. [15] [13] Also, olfaction and visual signaling at a distance of at least 20m. [16] Lorises interact throughout the night and sleep in groups during the daytime. Females with exclusive home ranges, rarely interact with other females except mothers and daughters. Females are more affiliated with males that are in the same sleeping area. Males tend to be more aggressive to other males that are not associated with their sleeping area. However, there is positive interaction with males that are within the sleeping area. The males also tend to infants that are in the sleeping area and sleeping arrangements are the only social behaviour lorises take part in.

From Kannur, Kerala, India at day. Grey Slender Loris imported from iNaturalist photo 52202458 on 12 March 2023.jpg
From Kannur, Kerala, India at day.

Reproduction

Lorises are found to be promiscuous, multiple males competing for one female for mating. Mating occurs biannually, April–June, and October–December. [13] Although mating occurs twice a year, lorises can only breed once a year. One estrus cycle will fail and another will begin approximately 2 weeks following. Gestation is approximately 170 days. Twins are very common among lorises. The reproductive tract and ovarian cycle reflect the general mammalian plans. As far as lactation, four teats develop and produce milk before the infant is born. After giving birth, females are non-receptive to males and care for their young. [17] The reproductive rate of the slender loris is among the lowest of any primate under 500g. [18]

Geographic range and habitat

Found in southern India and Sri Lanka, the gray slender loris inhabits primary and secondary rainforest, dry semi-deciduous forest, and montane cloud forest up to 2,000 metres (6,600 ft) above sea level. [11] It is found in south-western India roughly between the Tapti and Godavari Rivers down to the south coast of the subcontinent. The subspecies are separated geographically. In south-western India, the Malabar gray slender loris occurs in the wet forests of the Western Ghats in the states of Karnataka, Kerala and Tamil Nadu up to an altitude of 1,200 metres (3,900 ft). The Mysore gray slender loris inhabits the tropical dry forests of the Eastern Ghats in the states of Andhra Pradesh, Karnataka and Tamil Nadu, frequenting open Euphorbia scrub forests and Acacia trees at an altitude of 300–500 metres (980–1,640 ft). [19] [14] [20] [21] It can also be found on the dry eastern slopes of the Western Ghats. [21]

Conservation status

Although considered near threatened on the IUCN Red List and classified under Schedule I (Part 1) of the Indian Wildlife Act, 1972, the threat to these primates is increasing. Loris is used to make love potions, treat leprosy and eye ailments. Some villagers keep lorises as pets, but because they are difficult to maintain there is high mortality. Some cities have them captive in zoos, and astrologers use them to pick out tarot cards. [22] Insects form the main source of food for lorises. Farmlands, which are high in cattle waste, produce large amounts of insects. Thus lorises do not compete with humans for resources and therefore tend to be tolerated by them. [23] Habitat fragmentation is also a threat to the loris population, as well as loss of acacia trees, which is a preferred tree species for the loris. [20] Conservation efforts are developing and more research on conservation efforts are ongoing. [14]

Related Research Articles

<span class="mw-page-title-main">Lorisidae</span> Family of primates

Lorisidae is a family of strepsirrhine primates. The lorisids are all slim arboreal animals and comprise the lorises, pottos and angwantibos. Lorisids live in tropical, central Africa as well as in south and southeast Asia.

<span class="mw-page-title-main">Loris</span> Subfamily of primates

Loris is the common name for the strepsirrhine mammals of the subfamily Lorinae in the family Lorisidae. Loris is one genus in this subfamily and includes the slender lorises, Nycticebus is the genus containing the slow lorises, and Xanthonycticebus is the genus name of the pygmy slow loris.

<span class="mw-page-title-main">Red slender loris</span> Species of primate

The red slender loris is a small, nocturnal strepsirrhine primate native to the rainforests of Sri Lanka. This is No. 6 of the 10 focal species and No. 22 of the 100 EDGE mammal species worldwide considered the most evolutionarily distinct and globally endangered. Two subspecies have been identified, L. t. tardigradus and L. t. nycticeboides.

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

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. Traditionally only the 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.

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

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">Slow loris</span> Genus of primates from Southeast Asia

Slow lorises are a group of several species of nocturnal strepsirrhine primates that make up the genus Nycticebus. Found in Southeast Asia and bordering areas, they range from Bangladesh and Northeast India in the west to the Sulu Archipelago in the Philippines in the east, and from Yunnan province in China in the north to the island of Java in the south. Although many previous classifications recognized as few as a single all-inclusive species, there are now at least eight that are considered valid: the Sunda slow loris (N. coucang), Bengal slow loris (N. bengalensis), Javan slow loris (N. javanicus), Philippine slow loris (N. menagensis), Bangka slow loris (N. bancanus), Bornean slow loris (N. borneanus), Kayan River slow loris (N. kayan) and Sumatran slow loris. A ninth species, the pygmy slow loris (X. pygmaeus), was recently moved to the new genus Xanthonycticebus. After the pygmy slow loris, the group's closest relatives are the slender lorises of southern India and Sri Lanka. Their next closest relatives are the African lorisids, the pottos, false pottos, and angwantibos. They are less closely related to the remaining lorisoids, and more distantly to the lemurs of Madagascar. Their evolutionary history is uncertain since their fossil record is patchy and molecular clock studies have given inconsistent results.

<span class="mw-page-title-main">Sunda slow loris</span> Species of primate found in South East Asia

The Sunda slow loris, or greater slow loris, is a strepsirrhine primate and a species of slow loris native to Indonesia, West Malaysia, southern Thailand and Singapore. It measures 27 to 38 cm from head to tail and weighs between 599 and 685 g. Like other slow lorises, it has a wet nose (rhinarium), a round head, small ears hidden in thick fur, a flat face, large eyes and a vestigial tail.

<span class="mw-page-title-main">Pygmy slow loris</span> Species of primate

The pygmy slow loris is a species of slow loris found east of the Mekong River in Vietnam, Laos, eastern Cambodia, and China. It occurs in a variety of forest habitats, including tropical dry forests, semi-evergreen, and evergreen forests. It was originally classified within Nycticebus until it was transferred to the genus Xanthonycticebus in 2022. Two species are recognised, the northern pygmy loris X. intermedius from northern Vietnam, Laos and China and the southern pygmy loris X. pygmaeus from southern Vietnam, Laos and Cambodia. The animal is nocturnal and arboreal, crawling along branches using slow movements in search of prey. Unlike other primates, it does not leap. It lives together in small groups usually with one or two offspring. An adult can grow to around 19 to 23 cm long and has a very short tail. It weighs about 450 g (1.0 lb). Its diet consists of fruits, insects, small fauna, tree sap, and floral nectar. The animal has a toxic bite, which it gets by licking a toxic secretion from glands on the inside of its elbows. The teeth in its lower jaw form a comb-like structure called a toothcomb that is used for scraping resin from tree bark.

<span class="mw-page-title-main">Bengal slow loris</span> Species of primate

The Bengal slow loris or northern slow loris is a strepsirrhine primate and a species of slow loris native to the Indian subcontinent and Indochina. Its geographic range is larger than that of any other slow loris species. Considered a subspecies of the Sunda slow loris (N. coucang) until 2001, phylogenetic analysis suggests that the Bengal slow loris is most closely related to the Sunda slow loris. However, some individuals in both species have mitochondrial DNA sequences that resemble those of the other species, due to introgressive hybridization. It is the largest species of slow loris, measuring 26 to 38 cm from head to tail and weighing between 1 and 2.1 kg. Like other slow lorises, it has a wet nose (rhinarium), a round head, flat face, large eyes, small ears, a vestigial tail, and dense, woolly fur. The toxin it secretes from its brachial gland differs chemically from that of other slow loris species and may be used to communicate information about sex, age, health, and social status.

<span class="mw-page-title-main">Slender loris</span> Genus of primates

The slender lorises (Loris) are a genus of loris native to India and Sri Lanka. The genus comprises two species, the red slender loris found in Sri Lanka and the gray slender loris from Sri Lanka and India. Slender lorises spend most of their life in trees, traveling along the tops of branches with slow and precise movements. They are found in tropical rainforests, scrub forests, semi-deciduous forests, and swamps. The primates have lifespans of approximately 15 years and are nocturnal. Slender lorises generally feed on insects, reptiles, plant shoots, and fruit.

<span class="mw-page-title-main">Javan slow loris</span> Species of primate native to Indonesia

The Javan slow loris is a strepsirrhine primate and a species of slow loris native to the western and central portions of the island of Java, in Indonesia. Although originally described as a separate species, it was considered a subspecies of the Sunda slow loris (N. coucang) for many years, until reassessments of its morphology and genetics in the 2000s resulted in its promotion to full species status. It is most closely related to the Sunda slow loris and the Bengal slow loris (N. bengalensis). The species has two forms, based on hair length and, to a lesser extent, coloration.

<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">Philippine slow loris</span> Species of primate

The Philippine slow loris is a strepsirrhine primate and a species of slow loris that is native to the north and east coastal areas of the island of Borneo, as well as the Sulu Archipelago in the Philippines. The species was first named as the Bornean slow loris in 1892, but lumped into the widespread Sunda slow loris (N. coucang) in 1952. However, it was promoted to full species status – again as the Bornean slow loris – based on molecular analysis in 2006. In 2013, two former subspecies of the Bornean slow loris were elevated to species status, and a new species—N. kayan—was recognized among the Bornean population.

<span class="mw-page-title-main">William Charles Osman Hill</span> British anthropologist and anatomist (1901–1975)

William Charles Osman Hill FRSE FZS FLS FRAI was a British anatomist, primatologist, and a leading authority on primate anatomy during the 20th century. He is best known for his nearly completed eight-volume series, Primates: Comparative Anatomy and Taxonomy, which covered all living and extinct primates known at the time in full detail and contained illustrations created by his wife, Yvonne. Schooled at King Edward VI Camp Hill School for Boys in Birmingham and University of Birmingham, he went on to publish 248 works and accumulated a vast collection of primate specimens that are now stored at the Royal College of Surgeons of England.

<span class="mw-page-title-main">Conservation of slow lorises</span> Conservation management of the nocturnal primates in Asia

Slow lorises are nocturnal strepsirrhine primates in the genus Nycticebus that live in the rainforests of South and Southeast Asia. They are threatened by habitat loss and fragmentation from deforestation, selective logging, and slash-and-burn agriculture, as well as by collection and hunting for the wildlife trade, including the exotic pet trade, and for use in traditional medicine and as bushmeat. Because of these and other threats, all five species of slow loris are listed as either "Vulnerable" or "Endangered" by the International Union for Conservation of Nature (IUCN). Their conservation status was originally listed as "Least Concern" in 2000 because of imprecise population surveys and the frequency in which these primates were found in animal markets. Because of their rapidly declining populations and local extinctions, their status was updated and in 2007 the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES) elevated them to Appendix I, which prohibits international commercial trade. Local laws also protect slow lorises from hunting and trade, but enforcement is lacking in most areas.

<i>Nycticebus kayan</i> Species of primate

The Kayan River slow loris is a strepsirrhine primate and a species of slow loris that is native to the northern and central highland region of the island of Borneo. The species was originally thought to be a part of the Bornean slow loris (N. menagensis) population until 2013, when a study of museum specimens and photographs identified distinct facial markings, which helped to differentiate it. It is distinguished by the high contrast of its black and white facial features, as well as the shape and width of the stripes of its facial markings.

The Bangka slow loris is a strepsirrhine primate and a species of slow loris that is native to southwestern Borneo and the island of Bangka. Originally considered a subspecies or synonym of the Bornean slow loris (N. menagensis), it was promoted to full species status in 2013 when a study of museum specimens and photographs identified distinct facial markings, which helped to differentiate it as a separate species. It is distinguished by the crimson red fur on its back, light-colored facial features, as well as the shape and width of the stripes of its facial markings.

<i>Nycticebus borneanus</i> Species of primate

Nycticebus borneanus, the Bornean slow loris, is a strepsirrhine primate and a species of slow loris that is native to central south Borneo in Indonesia. Formerly considered a subspecies or synonym of N. menagensis, it was promoted to full species status in 2013 when a study of museum specimens and photographs identified distinct facial markings, which helped to differentiate it as a separate species. It is distinguished by its dark, contrasting facial features, as well as the shape and width of the stripes of its facial markings.

Anna Nekaris is a British anthropologist and professor in primate conservation in the School of Law and Social Sciences at Oxford Brookes University. She is interested in the conservation of Asian nocturnal animals and African nocturnal primates. She was appointed an Officer of the Order of the British Empire in the 2024 New Year Honours.

References

  1. 1 2 Dittus, W.; Singh, M.; Gamage, S.N.; Kumara, H.N.; Kumar, A.; Nekaris, K.A.I. (2020). "Loris lydekkerianus". IUCN Red List of Threatened Species . 2020: e.T44722A17970358. doi: 10.2305/IUCN.UK.2020-3.RLTS.T44722A17970358.en . Retrieved 17 November 2021.
  2. 1 2 3 Groves, C. P. (2005). Wilson, D. E.; Reeder, D. M. (eds.). Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Baltimore: Johns Hopkins University Press. p. 122. ISBN   0-801-88221-4. OCLC   62265494.
  3. ↑ Osman Hill, W. C. (1953). Primates Comparative Anatomy and Taxonomy I—Strepsirhini. Edinburgh Univ Pubs Science & Maths, No 3. Edinburgh University Press. OCLC   500576914.
  4. ↑ Groves, Colin (1998). "Systematics of tarsiers and lorises". Primates. 39: 13–27. doi:10.1007/BF02557740. S2CID   10869981.
  5. 1 2 Nekaris, K. A. I.; Jayawardene, J. (2004). "Survey of slender loris (Primates, Lorisidae Grey, 1821: Loris tardigradus Linnaeus, 1758 and Loris lydekkerianus Cabrera, 1908) in Sri Lanka". J. Zool. Lond. 262: 327–338. doi:10.1017/S0952836903004710.
  6. ↑ Nekaris, K. A. I.; Roos, C.; Pimley, E. R.; Schulze, H. (2006). "Diversity slowly coming to light: reconsidering the taxonomy of pottos and lorises". Int. J. Primatol. 27 (suppl. 1): 441.
  7. ↑ Gamage, S. N.; Groves, C. P.; Marikar, F. M. M. T.; Turner, C. S.; Padmalal, U. K. G. K.; Kotagama, S. W. (2017). "The taxonomy, distribution, and conservation status of the slender loris (Primates, Lorisidae: Loris) in Sri Lanka". Journal of Primate Conservation. 30: 1–25.
  8. ↑ Perera, M.; Sandun, J. (2008). "A Review of the Distribution of Grey Slender Loris (Loris lydekkerianus) in Sri Lanka" (PDF). Primate Conservation. 23: 89–96. doi:10.1896/052.023.0110. S2CID   8941145. Archived from the original (PDF) on 2009-02-05.
  9. ↑ Yapa, Asoka C. (2013). The Mammals of Sri Lanka. ISBN   9789558576328.
  10. ↑ Gamage, Saman; Marikar, Faiz; Groves, Colin; Turner, Craig; Padmalal, Kalinga; Kotagama, Sarath (2019). "Phylogenetic relationship among slender loris species (Primates, Lorisidae: Loris) in Sri Lanka based on mtDNA CO1 barcoding". Turkish Journal of Zoology. 43 (6): 609–616. doi: 10.3906/zoo-1901-20 .
  11. 1 2 Nekaris, K. A. I. (2013). "Family Lorisidae (Angwantibos, Pottos and Lorises)". In Mittermeier, R. A.; Rylands, A. B.; Wilson, D. E. (eds.). Handbook of the Mammals of the World. Vol. 3. Barcelona: Lynx Edicions. pp. 211–235. ISBN   978-84-96553-89-7.
  12. 1 2 Radhakrishna, S. (2010). "Behavioural variation in the Mysore slender loris Loris lydekkerianus lydekkerianus" (PDF). Current Science (Bangalore) (pdf). 99 (9): 1226–1232.
  13. 1 2 3 Radhakrishna, S.; Singh, M. (2002). "Social behaviour of the slender loris (Loris tardigradus lydekkerianus)". Folia Primatologica. 73 (4): 181–196. doi:10.1159/000065426. ISSN   0015-5713. PMID   12399658. S2CID   3013829.
  14. 1 2 3 Singh, Mewa; Lindburg, Donald G.; Udhayan, A.; Kumar, M. Anand; Kumara, H. N. (1999). "Status survey of slender loris Loris tardigradus lydekkerianus in Dindigul, Tamil Nadu, India". Oryx. 33: 31–37. doi: 10.1046/j.1365-3008.1999.00033.x .
  15. ↑ Radhakrishna, S.; Singh, M. (2002). "Home range and ranging pattern in the slender loris (Loris tardigradus lydekkerianus)". Primates. 43 (3): 237–248. doi:10.1007/BF02629651. PMID   12145404. S2CID   31980000.
  16. ↑ Nekaris, K.A.I. (2006). "Social lives of adult mysore slender lorises (Loris lydekkerianus lydekkerianus)". American Journal of Primatology. 68 (12): 1171–1182. doi:10.1002/ajp.20316. PMID   17096424. S2CID   17932047.
  17. ↑ Ramaswami, L. S.; Kumar, T. C. A. (1965). "Some aspects of reproduction of the female slender loris, Loris tardigradus lydekkerianus Cabr". Acta Zoologica. 46 (3): 257–273. doi:10.1111/j.1463-6395.1965.tb00734.x.
  18. ↑ Izard, M. Kay; Rasmussen, D. Tab (1985). "Reproduction in the slender loris (Loris tardigradus malabaricus)". American Journal of Primatology. 8 (2): 153–165. doi:10.1002/ajp.1350080206. PMID   31986817. S2CID   85156708.
  19. ↑ Roos, C.; Boonratana, R.; Supriatna, J.; Fellowes, J. R.; Groves, C. P.; Nash, S. D.; Rylands, A. B.; Mittermeier, R. A. (2014). "An updated taxonomy and conservation status review of Asian primates" (PDF). Asian Primates Journal (pdf). 4 (1): 1–38.
  20. 1 2 Kumara, H. N. (2006). "Distribution, habitat correlates, and conservation of Loris lydekkerianus in Karnataka, India" (PDF). International Journal of Primatology (pdf). 27 (4): 941–969. doi:10.1007/s10764-006-9054-z. S2CID   22320115.[ dead link ]
  21. 1 2 Radhakrishna, S.; Kumara, H. N.; Sasi, R. (2011). "Distribution patterns of slender loris subspecies (Loris lydekkerianus) in Kerala, Southern India". International Journal of Primatology. 32 (4): 1007–1019. doi:10.1007/s10764-011-9518-7. S2CID   31225684.
  22. ↑ Kanagavel, A.; Sinclair, C.; Sekar, R.; Raghavan, R. (2013). "Moolah, misfortune or spinsterhood? The plight of slender loris Loris lydekkerianus in southern India". Journal of Threatened Taxa. 5: 3585–3588. doi: 10.11609/jott.o3265.948 .
  23. ↑ Singh, M.; Kumar, M. A.; Kumara, H. N.; Mohnot, S. M. (2000). "Distribution and conservation of slender lorises in southern Andhra Pradesh, South India". International Journal of Primatology. 21 (4): 721–730. doi:10.1023/A:1005573506489. S2CID   33800284.
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