Nycticebus Kayan
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The Kayan River slow loris (''Nycticebus kayan'') is a
strepsirrhine Strepsirrhini or Strepsirhini (; ) is a suborder of primates that includes the lemuriform primates, which consist of the lemurs of Madagascar, galagos ("bushbabies") and pottos from Africa, and the lorises from India and southeast Asia. Colle ...
primate Primates are a diverse order of mammals. They are divided into the strepsirrhines, which include the lemurs, galagos, and lorisids, and the haplorhines, which include the tarsiers and the simians ( monkeys and apes, the latter including ...
and a
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
of
slow loris 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 Archip ...
that is native to the northern and central highland region of the island of
Borneo Borneo (; id, Kalimantan) is the third-largest island in the world and the largest in Asia. At the geographic centre of Maritime Southeast Asia, in relation to major Indonesian islands, it is located north of Java, west of Sulawesi, and ea ...
. 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 species is named after the
Kayan River The Kayan River is a river of Borneo island, flowing in the North Kalimantan province of Indonesia, about 1600 km northeast of the capital Jakarta. Tributaries include the Bahau River. Hydrology The Kayan River rises on Mount Ukeng, pa ...
, which runs through its native habitat. As with other slow lorises, this
arboreal Arboreal locomotion is the locomotion of animals in trees. In habitats in which trees are present, animals have evolved to move in them. Some animals may scale trees only occasionally, but others are exclusively arboreal. The habitats pose nu ...
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 The International Union for Conservation of Nature (IUCN; officially International Union for Conservation of Nature and Natural Resources) is an international organization working in the field of nature conservation and sustainable use of nat ...
(IUCN), it is likely to be listed as " Vulnerable" or placed in a higher-risk category when its
conservation status The conservation status of a group of organisms (for instance, a species) indicates whether the group still exists and how likely the group is to become extinct in the near future. Many factors are taken into account when assessing conservatio ...
is assessed. It is primarily threatened by
habitat loss Habitat destruction (also termed habitat loss and habitat reduction) is the process by which a natural habitat becomes incapable of supporting its native species. The organisms that previously inhabited the site are displaced or dead, thereby ...
and the illegal
wildlife trade Wildlife trade refers to the of products that are derived from non-domesticated animals or plants usually extracted from their natural environment or raised under controlled conditions. It can involve the trade of living or dead individuals, ti ...
.


Taxonomy and phylogeny

''N. kayan'' is a strepsirrhine primate and a species of slow loris (known collectively as the
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
''Nycticebus''), within the loris
family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...
(
Lorisidae Lorisidae (or sometimes Loridae) 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 As ...
). Prior to 2013, museum specimens of this animal had been identified as the Bornean slow loris (''Nycticebus menagensis''), which had first been described by the English naturalist
Richard Lydekker Richard Lydekker (; 25 July 1849 – 16 April 1915) was an English naturalist, geologist and writer of numerous books on natural history. Biography Richard Lydekker was born at Tavistock Square in London. His father was Gerard Wolfe Lydekker, ...
in 1893 as ''Lemur menagensis''. However, in 1939 Reginald Innes Pocock consolidated all slow lorises into a single
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
, ''N. coucang''. Primatologist
William Charles Osman Hill Dr William Charles Osman Hill FRSE FZS FLS FRAI (13 July 1901 – 25 January 1975) 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-vol ...
continued this course in his influential 1953 book, ''Primates: Comparative Anatomy and Taxonomy''. In 1971, that view was refined by
Colin Groves Colin Peter Groves (24 June 1942 – 30 November 2017) was a British-Australian biologist and anthropologist. Groves was Professor of Biological Anthropology at the Australian National University in Canberra, Australia. Education Born in Englan ...
, who recognized the
pygmy slow loris The pygmy slow loris (''Xanthonycticebus pygmaeus'') 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-everg ...
(''N. pygmaeus'') as a separate species. He also divided ''N. coucang'' into four subspecies, – one of those being ''N. coucang menagensis'', the Bornean slow loris. The subspecies ''N. c. menagensis'' was elevated to the species level (as ''N. menagensis'') in 2006, when molecular analysis showed it to be genetically distinct from ''N. coucang''. A 2013 review of all museum specimens and photographs attributed to ''N. menagensis'' showed that they in fact comprised a
species complex In biology, a species complex is a group of closely related organisms that are so similar in appearance and other features that the boundaries between them are often unclear. The taxa in the complex may be able to hybridize readily with each oth ...
. First, two former subspecies of ''N. menagensis'' were elevated to the two distinct species '' N. bancanus'' and '' N. borneanus''. Further, ''N. kayan'' was recognized as a new species that is also distinct from the nominate subspecies, ''N. menagensis''. All newly recognized or elevated species showed significant differences in their "facemask"—the coloration patterns on their face. Analysis of the facemask patterns suggests that ''N. kayan'' diverged from ''N. menagensis'' and ''N. borneanus'' through sympatric speciation (divergent evolution of organisms living in the same geographic region), while geographic barriers may account for its divergence with ''N. bancanus'' (
allopatric speciation Allopatric speciation () – also referred to as geographic speciation, vicariant speciation, or its earlier name the dumbbell model – is a mode of speciation that occurs when biological populations become geographically isolated from ...
). ''N. kayan'' is named after the
Kayan River The Kayan River is a river of Borneo island, flowing in the North Kalimantan province of Indonesia, about 1600 km northeast of the capital Jakarta. Tributaries include the Bahau River. Hydrology The Kayan River rises on Mount Ukeng, pa ...
, which runs through its native habitat and near Peleben, the type locality of the original specimen. The
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of sever ...
, AMNH 106012, was originally collected on 8 October 1935 by Baron V. von Plessen near Peleben in the province of
East Kalimantan East Kalimantan ( Indonesian: ) is a province of Indonesia. Its territory comprises the eastern portion of Borneo. It had a population of about 3.03 million at the 2010 census (within the current boundary), 3.42 million at the 2015 census, and 3 ...
in
Borneo Borneo (; id, Kalimantan) is the third-largest island in the world and the largest in Asia. At the geographic centre of Maritime Southeast Asia, in relation to major Indonesian islands, it is located north of Java, west of Sulawesi, and ea ...
and is housed in the American Museum of Natural History in New York. It consists of a male skin and skull, with a head-body length of .


Physical description

Like other slow lorises, it has a
vestigial Vestigiality is the retention, during the process of evolution, of genetically determined structures or attributes that have lost some or all of the ancestral function in a given species. Assessment of the vestigiality must generally rely on co ...
tail, round head, and short ears. It has a
rhinarium The rhinarium (New Latin, "belonging to the nose"; plural: rhinaria) is the furless skin surface surrounding the external openings of the nostrils in many mammals. Commonly it is referred to as the tip of the ''snout'', and breeders of cats and ...
(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 Incisors (from Latin ''incidere'', "to cut") are the front teeth present in most mammals. They are located in the premaxilla above and on the mandible below. Humans have a total of eight (two on each side, top and bottom). Opossums have 18, wher ...
, 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 A grooming claw (or toilet claw) is the specialized claw or nail on the foot of certain primates, used for personal grooming. All prosimians have a grooming claw, but the digit that is specialized in this manner varies. Tarsiers have a grooming ...
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 A toothcomb (also tooth comb or dental comb) is a dental structure found in some mammals, comprising a group of front teeth arranged in a manner that facilitates grooming, similar to a hair comb. The toothcomb occurs in lemuriform primates ( ...
, 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 face mask of ''N. kayan'' differs from those of other Bornean lorises in several ways. First, the top of the dark ring around its eyes is either rounded or pointed (not diffuse at the edges) and the bottom stretches below the
zygomatic arch In anatomy, the zygomatic arch, or cheek bone, is a part of the skull formed by the zygomatic process of the temporal bone (a bone extending forward from the side of the skull, over the opening of the ear) and the temporal process of the zygo ...
, and sometimes extends as far down as the jaw. Second, the stripe between the eyes is occasionally bulb-shaped, compared to the rectangular stripe seen in the neighboring species. Also, a light band of fur in front of the ears is usually intermediate in width compared to the narrow and wide bands seen in the other Bornean species. Compared to ''N. menagensis'', the facemask of ''N. kayan'' has more contrast between its dark black and white features, and its ears are always covered in hair, whereas those of ''N. menagensis'' are generally naked. Overall, its fur is generally longer and fluffier compared to ''N. menagensis''. Based on a limited number of specimens, the species is about long, and weighs about .


Distribution

''N. kayan'' is found in central and northern Borneo. Its range extends south to the
Mahakam The Mahakam River ( Indonesian: ''Sungai Mahakam'') is third longest and volume discharge river in Borneo after Kapuas River and Barito River, it is located in Kalimantan, Indonesia. It flows from the district of Long Apari in the highlands ...
and
Rajang River The Rajang River ( ms, Batang Rajang) is a river in Sarawak, northwest Borneo, Malaysia. The river originates in the Iran Mountains, flows through Kapit, and then towards the South China Sea. At approximately , the river is the seventh-longest i ...
s in the
Indonesia Indonesia, officially the Republic of Indonesia, is a country in Southeast Asia and Oceania between the Indian and Pacific oceans. It consists of over 17,000 islands, including Sumatra, Java, Sulawesi, and parts of Borneo and New Guine ...
n province of East Kalimantan and the
Malaysia Malaysia ( ; ) is a country in Southeast Asia. The federation, federal constitutional monarchy consists of States and federal territories of Malaysia, thirteen states and three federal territories, separated by the South China Sea into two r ...
n province of
Sarawak Sarawak (; ) is a state of Malaysia. The largest among the 13 states, with an area almost equal to that of Peninsular Malaysia, Sarawak is located in northwest Borneo Island, and is bordered by the Malaysian state of Sabah to the northeast, ...
, respectively, and north to southern side of
Mount Kinabalu Mount Kinabalu ( ms, Gunung Kinabalu, Dusun: ''Gayo Ngaran or Nulu Nabalu'') is the highest mountain in Borneo and Malaysia. With an elevation of , it is third-highest peak of an island on Earth, and 20th most prominent mountain in the worl ...
in the Malaysian province of
Sabah Sabah () is a state of Malaysia located in northern Borneo, in the region of East Malaysia. Sabah borders the Malaysian state of Sarawak to the southwest and the North Kalimantan province of Indonesia to the south. The Federal Territory o ...
. Although it is not found along the coast, its range spans Borneo from east to west. Its range overlaps that of ''N. menagensis'' in East Kalimantan and Sabah, and ''N. borneanus'' is a neighboring species.


Habitat and ecology

Like other slow lorises, ''N. kayan'' is
arboreal Arboreal locomotion is the locomotion of animals in trees. In habitats in which trees are present, animals have evolved to move in them. Some animals may scale trees only occasionally, but others are exclusively arboreal. The habitats pose nu ...
, nocturnal, and
omnivorous An omnivore () is an animal that has the ability to eat and survive on both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize the nut ...
, 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 A toxin is a naturally occurring organic poison produced by metabolic activities of living cells or organisms. Toxins occur especially as a protein or conjugated protein. The term toxin was first used by organic chemist Ludwig Brieger (1849 ...
is produced by licking a brachial gland (a gland by its elbow), and the secretion mixes with its saliva to activate. The toxic bite is a deterrent to predators, and the toxin is also applied to the fur during grooming as a form of protection for infants. When threatened, slow lorises may also lick their brachial glands and bite the aggressors, delivering the toxin into the wound. 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 ''Nycticebus kayan'' 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 least, with some of them likely to be assigned to a higher-risk category. Between 1987 and 2012, one-third of Borneo's forests were lost, making habitat loss one of the greatest threats to the survival of ''N. kayan''. The illegal
wildlife trade Wildlife trade refers to the of products that are derived from non-domesticated animals or plants usually extracted from their natural environment or raised under controlled conditions. It can involve the trade of living or dead individuals, ti ...
is also a major factor, with loris parts commonly sold for
traditional medicine Traditional medicine (also known as indigenous medicine or folk medicine) comprises medical aspects of traditional knowledge that developed over generations within the folk beliefs of various societies, including indigenous peoples, before the ...
. Further,
viral video A viral video is a video that becomes popular through a viral process of Internet sharing, typically through video sharing websites such as YouTube as well as social media and email.Lu Jiang, Yajie Miao, Yi Yang, ZhenZhong Lan, Alexander Haupt ...
s on
YouTube YouTube is a global online video sharing and social media platform headquartered in San Bruno, California. It was launched on February 14, 2005, by Steve Chen, Chad Hurley, and Jawed Karim. It is owned by Google, and is the second mo ...
promote the
exotic pet An exotic pet is a pet which is relatively rare or unusual to keep, or is generally thought of as a wild species rather than as a domesticated pet. The definition varies by culture, location, and over time—as animals become firmly enough est ...
trade. However, all slow loris species are protected from commercial trade under Appendix I of the Convention on International Trade in Endangered Species of Wild Fauna and Flora (
CITES CITES (shorter name for the Convention on International Trade in Endangered Species of Wild Fauna and Flora, also known as the Washington Convention) is a multilateral treaty to protect endangered plants and animals from the threats of interna ...
).


References


Literature cited

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