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Nycticebus borneanus
Conservation status
Vulnerable  (IUCN 3.1)
CITES Appendix I (CITES)
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Order:	Primates
Suborder:	Strepsirrhini
Family:	Lorisidae
Genus:	Nycticebus
Species:	N. borneanus
Binomial name
Nycticebus borneanus (Lyon, 1906)

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.

As with other slow lorises, this arboreal and nocturnal species primarily eats insects, tree gum, nectar, and fruit and has a toxic bite, a unique feature among primates. Although not yet evaluated by the International Union for Conservation of Nature (IUCN), it is likely to be listed as "Vulnerable" or placed in a higher-risk category when its conservation status is assessed. It is primarily threatened by habitat loss and the illegal wildlife trade.

Taxonomy and phylogeny

N. borneanus is a strepsirrhine primate, and species of slow loris (genus Nycticebus) within the family Lorisidae. Museum specimens of this animal had previously been identified as the Bornean slow loris using the scientific name Nycticebus menagensis – first described by the English naturalist Richard Lydekker in 1893 as Lemur menagensis, – a scientific name now assigned exclusively to the Philippine slow loris. In 1906, Marcus Ward Lyon Jr. first described N. borneanus from western Borneo. By 1953, all of the slow lorises were lumped together into a single species, the Sunda slow loris (Nycticebus coucang). In 1971, that view was updated by distinguishing the pygmy slow loris (N. pygmaeus) as a species, and by further recognizing four subspecies, including N. coucang menagensis. From then until 2005, N. borneanus was considered a synonym of N. menagensis. The latter was elevated to the species level in 2006, when molecular analysis showed it to be genetically distinct from N. coucang.

A 2013 review of museum specimens and photographs attributed to N. menagensis resulted in elevating two of its former subspecies to species: N. bancanus and N. borneanus. Additionally, N. kayan emerged as a new species, which had previously been overlooked. All newly recognized or elevated species showed significant differences in their "face mask"—the coloration patterns on their face.

Physical description

Like other slow lorises, it has a vestigial tail, round head, and short ears. It has a rhinarium (the moist, naked surface around the nostrils of the nose) and a broad, flat face with large eyes. Like N. menagensis, this and all other Bornean species lack a second upper incisor, which distinguishes them from other slow lorises. On its front feet, the second digit is smaller than the rest; the big toe on its hind foot opposes the other toes, which enhances its gripping power. Its second toe on the hind foot has a curved grooming claw that it uses for scratching and grooming, while the other nails are straight. It also possesses a specialized arrangement of lower front teeth, called a toothcomb, which is also used for grooming, as with other lemuriform primates. On the ventral side of its elbow, it has a small swelling called the brachial gland, which secretes a pungent, clear oily toxin that the animal uses defensively by wiping it on its toothcomb.

The facial markings of N. borneanus are dark and contrasting. The dark rings around its eyes are usually rounded on top, though sometimes diffuse-edged, and they never reach below the zygomatic arch. The stripe between the eyes often varies in width, the ears are covered in hair, and the band of hair in front of the ears is wide. The colored patch on the top of the head is usually round, but is sometimes a narrower band. The body length averages 260.1 mm (10.24 in) for the species.

Distribution

N. borneanus is found in central south Borneo, in the Indonesian provinces of West, South, and Central Kalimantan. Its range extends south of the Kapuas River and east to the Barito River. However, N. borneanus is not found in the extreme southwest of the island. It may be sympatric with N. bancanus in the province of West Kalimantan.

Habitat and ecology

Like other slow lorises, N. borneanus is arboreal, nocturnal, and omnivorous, eating primarily insects, tree gum, nectar, and fruit. Likewise, this species has a toxic bite, a unique feature found only in slow lorises among primates. The toxin is produced by licking a brachial gland (a gland by their elbow), and the secretion mixes with its saliva to activate. Their toxic bite is a deterrent to predators, and the toxin is also applied to the fur during grooming as a form of protection for their infants. When threatened, slow lorises may also lick their brachial glands and bite their aggressors, delivering the toxin into the wounds. Slow lorises can be reluctant to release their bite, which is likely to maximize the transfer of toxins.

The face mask may help the species identify potential mates by distinguishing species, and may serve as an anti-predator strategy by making its eyes appear larger than they really are.

Conservation

While this new species has yet to be assessed by the IUCN, N. menagensis was listed as "Vulnerable" as of 2012. Because that species has been divided into four distinct species, each of the new species faces a higher risk of extinction. Accordingly, each of them are expected to be listed as "Vulnerable" at the least, with some of them likely to be assigned to a higher-risk category.

Between 1987 and 2012, one-third of Borneo's forests have been lost, making habitat loss one of the greatest threats to the survival of N. borneanus. The illegal wildlife trade is also a major factor, with loris parts commonly sold in traditional medicine and viral videos on YouTube promoting the exotic pet trade. However, all slow loris species are protected from commercial trade under Appendix I of CITES.

References

1. Nekaris, K.A.I.; Miard, P. (2020). "Nycticebus borneanus". IUCN Red List of Threatened Species. 2020. IUCN: e.T163015906A163015915. doi:10.2305/IUCN.UK.2020-2.RLTS.T163015906A163015915.en.
2. "Appendices I, II and III" (PDF). Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES). 2010. Archived from the original (PDF) on 2010-12-24. Retrieved 2012-12-14.
3. "Nycticebus borneanus". Integrated Taxonomic Information System. Retrieved 28 January 2016.
4. Munds, Nekaris & Ford 2013, p. 46.
5. Lyon 1906, pp. 535–536.
6. Osman Hill 1953, pp. 156–163.
7. Groves 1971.
8. Groves 2001, p. 99.
9. Groves 2005, p. 122.
10. Chen et al. 2006, p. 1198.
11. ^ Munds, Nekaris & Ford 2013, p. 47.
12. ^ Ankel-Simons 2007, p. 82.
13. ^ Smith & Xie 2008, pp. 159–160.
14. ^ Munds, Nekaris & Ford 2013, p. 53.
15. Ankel-Simons 2007, p. 246.
16. Hagey, Fry & Fitch-Snyder 2007, p. 253.
17. Munds, Nekaris & Ford 2013, p. 52–53.
18. Nekaris & Bearder 2007, pp. 28–33.
19. Alterman 1995, pp. 421–423.
20. Munds, Nekaris & Ford 2013, p. 49.
21. ^ Wall, T. (13 December 2012). "Three new species of venomous primate identified by MU researcher". Missouri University News Bureau. Archived from the original on 17 December 2012. Retrieved 19 December 2012.
22. Bryner, J. (14 December 2012). "Slow loris species, Nycticebus kayan, discovered in Borneo". The Huffington Post. Archived from the original on 17 December 2012. Retrieved 15 December 2012.
23. Walker, M. (13 December 2012). "Primate species: new slow loris found in Borneo". BBC News. Archived from the original on 25 December 2012.
24. Nekaris & Munds 2010, p. 390.

Literature cited

• Alterman, L. (1995). "Toxins and toothcombs: potential allospecific chemical defenses in Nycticebus and Perodicticus". In Alterman, L.; Doyle, G.A.; Izard, M.K (eds.). Creatures of the Dark: The Nocturnal Prosimians. New York, New York: Plenum Press. pp. 413–424. ISBN 978-0-306-45183-6. OCLC 33441731.
• Ankel-Simons, F. (2007). Primate Anatomy (3rd ed.). Academic Press. ISBN 978-0-12-372576-9.
• Chen, J. -H.; Pan, D.; Groves, C. P.; Wang, Y. -X.; Narushima, E.; Fitch-Snyder, H.; Crow, P.; Thanh, V. N.; Ryder, O.; Zhang, H. -W.; Fu, Y.; Zhang, Y. (2006). "Molecular phylogeny of Nycticebus inferred from mitochondrial genes". International Journal of Primatology. 27 (4): 1187–1200. doi:10.1007/s10764-006-9032-5. S2CID 24319996.
• Groves, Colin P. (1971). "Systematics of the genus Nycticebus" (PDF). Proceedings of the Third International Congress of Primatology. 1. Zürich, Switzerland: 44–53. Archived from the original (PDF) on 2011-02-28. Retrieved 2013-01-04.
• Groves, Colin P. (2001). Primate Taxonomy. Washington, DC: Smithsonian Institution Press. ISBN 978-1-56098-872-4.
• Groves, C. P. (2005). "Nycticebus menagensis". In Wilson, D. E.; Reeder, D. M (eds.). Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Johns Hopkins University Press. pp. 111–184. ISBN 978-0-8018-8221-0. OCLC 62265494.
• Hagey, L.R.; Fry, B.G.; Fitch-Snyder, H. (2007). "Talking defensively, a dual use for the brachial gland exudate of slow and pygmy lorises". In Gursky, S.L.; Nekaris, K.A.I. (eds.). Primate Anti-Predator Strategies. Developments in Primatology: Progress and Prospects. Springer. pp. 253–273. doi:10.1007/978-0-387-34810-0. ISBN 978-0-387-34807-0.
• Lyon, M.W. Jr. (1906). "Notes on the slow lemurs". Proceedings of the United States National Museum. 31 (1494): 527–538. doi:10.5479/si.00963801.31-1494.527.
• Munds, R. A.; Nekaris, K. A. I.; Ford, S. M. (2013) . "Taxonomy of the Bornean slow loris, with new species Nycticebus kayan (Primates, Lorisidae)" (PDF). American Journal of Primatology. 75 (1): 46–56. doi:10.1002/ajp.22071. PMID 23255350. S2CID 17077282.
• Nekaris, K.A.I.; Bearder, S.K. (2007). "Chapter 3: The Lorisiform Primates of Asia and Mainland Africa: Diversity Shrouded in Darkness". In Campbell, C.; Fuentes, C.A.; MacKinnon, K.; Panger, M.; Stumpf, R. (eds.). Primates in Perspective. New York, New York: Oxford University Press. pp. 28–33. ISBN 978-0-19-517133-4.
• Nekaris, K.A.I.; Munds, R. (2010). "Chapter 22: Using facial markings to unmask diversity: the slow lorises (Primates: Lorisidae: Nycticebus spp.) of Indonesia". In Gursky-Doyen, S.; Supriatna, J (eds.). Indonesian Primates. New York: Springer. pp. 383–396. doi:10.1007/978-1-4419-1560-3_22. ISBN 978-1-4419-1559-7.
• Osman Hill, W.C. (1953). Primates Comparative Anatomy and Taxonomy I—Strepsirhini. Edinburgh Univ Pubs Science & Maths, No 3. Edinburgh University Press. OCLC 500576914.
• Smith, Andrew T.; Xie, Yan (2008). A Guide to the Mammals of China. Princeton University Press. ISBN 978-0-691-09984-2.

• IUCN Red List vulnerable species
• Slow lorises
• Primates of Borneo
• Mammals described in 1906
• Taxa named by Marcus Ward Lyon Jr.
• Fauna of the Borneo lowland rain forests