Bornean Slow Loris
Encyclopedia
The Bornean slow loris (Nycticebus menagensis) or Philippine slow loris is a strepsirrhine
Strepsirrhini
The clade Strepsirrhini is one of the two suborders of primates. Madagascar's only non-human primates are strepsirrhines, and others can be found in southeast Asia and Africa...

 primate
Primate
A primate is a mammal of the order Primates , which contains prosimians and simians. Primates arose from ancestors that lived in the trees of tropical forests; many primate characteristics represent adaptations to life in this challenging three-dimensional environment...

 and a species
Species
In biology, a species is one of the basic units of biological classification and a taxonomic rank. A species is often defined as a group of organisms capable of interbreeding and producing fertile offspring. While in many cases this definition is adequate, more precise or differing measures are...

 of slow loris
Slow loris
Slow lorises are a group of five species of strepsirrhine primates which make up the genus Nycticebus. Found in South and Southeast Asia, they range from Northeast India in the west to the Philippines in the east, and from the Yunnan province in China in the north to the island of Java in the south...

 that is native to the island of Borneo
Borneo
Borneo is the third largest island in the world and is located north of Java Island, Indonesia, at the geographic centre of Maritime Southeast Asia....

 (including the Malaysian, Indonesian, and Bruneian portions), the nearby islands of Belitung
Belitung
Belitung, , is an island on the east coast of Sumatra, Indonesia in the Java Sea. The island is known for its pepper and for its tin. It was in the possession of the British from 1812 until the British ceded control of the island to the Dutch in the Anglo-Dutch Treaty of 1824...

 and Bangka
Bangka Island
Bangka is an island lying east of Sumatra, Indonesia. Population 626,955. Area: c.4,600 sq mi .There is an additional small island named Pulau Bangka in northern Sulawesi, Indonesia.-Geography:...

 in Indonesia
Indonesia
Indonesia , officially the Republic of Indonesia , is a country in Southeast Asia and Oceania. Indonesia is an archipelago comprising approximately 13,000 islands. It has 33 provinces with over 238 million people, and is the world's fourth most populous country. Indonesia is a republic, with an...

, and the Sulu Archipelago
Sulu Archipelago
The Sulu Archipelago is a chain of islands in the southwestern Philippines. This archipelago is considered to be part of the Moroland by the local rebel independence movement. This island group forms the northern limit of the Celebes Sea....

 in the Philippines
Philippines
The Philippines , officially known as the Republic of the Philippines , is a country in Southeast Asia in the western Pacific Ocean. To its north across the Luzon Strait lies Taiwan. West across the South China Sea sits Vietnam...

. The species was first named in 1892, but lumped into the widespread Sunda slow loris (N. coucang) in 1952. However, it was promoted to full species status based on molecular analysis in 2006.

Weighing 265 –, it is one of the smallest of the slow lorises, and can be distinguished from other slow lorises by its pale golden to red fur, the lack of markings on its head, and consistent absence of a second upper incisor
Incisor
Incisors are the first kind of tooth in heterodont mammals. They are located in the premaxilla above and mandible below.-Function:...

. Like other slow lorises, it has a vestigial tail, round head, short ears, a curved toilet-claw
Toilet-claw
A toilet-claw is the specialized claw or nail on the foot of certain primates, used for personal grooming. All prosimians have a toilet claw, but the digit that is specialized in this manner varies. Tarsiers have a toilet claw on toe two and toe three...

 for grooming, and a gland that produces an oily toxin that the animal uses for defense. The Bornean slow loris is arboreal, nocturnal, and occurs in low densities
Population density
Population density is a measurement of population per unit area or unit volume. It is frequently applied to living organisms, and particularly to humans...

, making it difficult to locate. It is also the least studied of Indonesia's slow lorises. It is found at elevations between 35 – in primary and secondary
Secondary forest
A secondary forest is a forest or woodland area which has re-grown after a major disturbance such as fire, insect infestation, timber harvest or windthrow, until a long enough period has passed so that the effects of the disturbance are no longer evident...

 lowland forest, gardens, and plantations. Information about its diet is limited, but it is suspected to be one of the more insectivorous
Insectivore
An insectivore is a type of carnivore with a diet that consists chiefly of insects and similar small creatures. An alternate term is entomophage, which also refers to the human practice of eating insects....

 slow loris species, and is also known to eat gum
Gum (botany)
Gum is a sap or other resinous material associated with certain species of the plant kingdom. This material is often polysaccharide-based and most frequently is associated with woody plants, particularly under the bark or as a seed coating...

 from woody plants.

The Bornean slow loris is classified as "Vulnerable
Vulnerable species
On 30 January 2010, the IUCN Red List of Threatened Species identified 9694 Vulnerable species, subspecies and varieties, stocks and sub-populations.-References:...

" by the International Union for Conservation of Nature (IUCN), is included in CITES Appendix I, which prevents international commercial trade, and is protected by Indonesian law. The species is found in numerous protected area
Protected area
Protected areas are locations which receive protection because of their recognised natural, ecological and/or cultural values. There are several kinds of protected areas, which vary by level of protection depending on the enabling laws of each country or the regulations of the international...

s within its range, making it the least threatened of the slow lorises. It is sparsely distributed throughout its range and is threatened by the illegal exotic pet
Exotic pet
An exotic pet is a rare or unusual animal pet, or an animal kept as a pet which is not commonly thought of as a pet.The definition is an evolving one; some rodents, reptiles, and amphibians have become firmly enough established in the world of animal fancy to no longer be considered exotic...

 trade and habitat loss.

Taxonomy and phylogeny

The Bornean slow loris was first described based on specimens collected by Frank S. Bourns and Dean C. Worcester during the Menage Scientific Expedition to the Philippines and Borneo in the early 1890s. The original collection was made between 5 October and 5 November 1891 in Tataan, Tawi-Tawi
Tawi-Tawi
Tawi-Tawi is an island province of the Philippines located in the Autonomous Region in Muslim Mindanao . The capital of Tawi-Tawi is Bongao. The province is the southernmost of the country sharing sea borders with the Malaysian State of Sabah and the Indonesian East Kalimantan province. To the...

 Island, in the Philippines. The specimens were given to Henry F. Nachtrieb, President of the Minnesota Academy of Sciences and Chairman of the Zoology Department at the University of Minnesota
University of Minnesota
The University of Minnesota, Twin Cities is a public research university located in Minneapolis and St. Paul, Minnesota, United States. It is the oldest and largest part of the University of Minnesota system and has the fourth-largest main campus student body in the United States, with 52,557...

. Nachtrieb was the first to use the name menagensis in 1892, based on a description of the species sent to him by Worcester in 1891. Worcester's progress report included a description of the specimen and an explanation on how they had obtained it:

Nachtrieb did not assign the name to a specific genus, noting that it was "an undescribed member of the Lemuridae". The following year, the English naturalist Richard Lydekker
Richard Lydekker
Richard Lydekker was an English naturalist, geologist and writer of numerous books on natural history.-Biography:...

 published the combination Lemur menagensis in the Zoological Record. This makes Lydekker the authority of the species name menagensis, because he was the first to use the specific name in combination with the name of a genus, although some subsequent authors credited other workers.

In his influential 1952 book Primates: Comparative Anatomy and Taxonomy, the primatologist William Charles Osman Hill
William Charles Osman Hill
William Charles Osman Hill was a British anatomist, primatologist, and a leading authority on primate anatomy during the 20th century...

 placed all the slow lorises in one species
Species
In biology, a species is one of the basic units of biological classification and a taxonomic rank. A species is often defined as a group of organisms capable of interbreeding and producing fertile offspring. While in many cases this definition is adequate, more precise or differing measures are...

, N. coucang. In 1971, however, Colin Groves
Colin Groves
Colin Peter Groves is Professor of Biological Anthropology at the Australian National University in Canberra, Australia.Born in England on 24 June 1942, Colin Groves completed a Bachelor of Science at University College London in 1963, and a Doctor of Philosophy at the Royal Free Hospital School of...

 recognized the pygmy slow loris
Pygmy Slow Loris
The pygmy slow loris is a rare species of loris found in the tropical dry forests of Vietnam, Laos, southern China, and east of the Mekong River in Cambodia....

 (N. pygmaeus) as a separate species, and divided N. coucang into four subspecies, including Nycticebus coucang menagensis. The Bornean slow loris was elevated to the species level (as Nycticebus menagensis) in 2006 when molecular analysis of DNA sequence
DNA sequence
The sequence or primary structure of a nucleic acid is the composition of atoms that make up the nucleic acid and the chemical bonds that bond those atoms. Because nucleic acids, such as DNA and RNA, are unbranched polymers, this specification is equivalent to specifying the sequence of...

s of the D-loop
D-loop
In molecular biology, a displacement loop or D-loop is a DNA structure where the two strands of a double-stranded DNA molecule are separated for a stretch and held apart by a third strand of DNA. The third strand has a base sequence which is complementary to one of the main strands and pairs with...

 and the cytochrome b
Cytochrome b
Cytochrome b/b6 is the main subunit of transmembrane cytochrome bc1 and b6f complexes. In addition, it commonly refers to a region of mtDNA used for population genetics and phylogenetics.- Function :...

 gene demonstrated it to be genetically distinct from N. coucang. The genetic evidence was corroborated by both a previous study (1998) on morphology
Morphology (biology)
In biology, morphology is a branch of bioscience dealing with the study of the form and structure of organisms and their specific structural features....

 (based on craniodental measurements) that indicated distinct differences between the subspecies that were consistent with separation at the species level, and a later study (2010) of facial markings.

Physical description

The body weight of the Bornean slow loris is typically in the range of 265 –, although weights of up to 700 grams (24.7 oz) have been recorded. One of the smallest of the slow lorises, it can be distinguished from the other species by its pale golden to red fur, light marks on its head, and the consistent lack of a second upper incisor
Incisor
Incisors are the first kind of tooth in heterodont mammals. They are located in the premaxilla above and mandible below.-Function:...

. Its skull length ranges between 54.5 and 56.5 mm (2.1 and 2.2 in), roughly intermediate in size between the smaller pygmy slow loris and the larger Sunda slow loris. The Bornean slow loris always has patches encircling the eye that end just below it. In contrast, the Sunda slow loris is characterized by medium width hair anterior to the opening of the ear, and the Javan slow loris always has a diamond stripe between the eyes.

Like other slow lorises, its tail is vestigial and it has a round head and short ears. It has a rhinarium
Rhinarium
The rhinarium is the moist, naked surface around the nostrils of the nose in most mammals. In actual scientific usage it is typically called a "wet snout" or "wet nose" from its moist and shiny appearance...

 (the moist, naked surface around the nostrils of the nose) and a broad, flat face with large eyes. 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 "toilet-claw
Toilet-claw
A toilet-claw is the specialized claw or nail on the foot of certain primates, used for personal grooming. All prosimians have a toilet claw, but the digit that is specialized in this manner varies. Tarsiers have a toilet claw on toe two and toe three...

" that the animal uses for scratching and grooming, while the other nails are straight. It also has a small swelling on the ventral side of its elbow called the brachial gland, which secretes a pungent, clear oily toxin that the animal uses defensively by wiping it on its toothcomb
Toothcomb
A toothcomb is an anatomical structure found in strepsirrhine primates, which includes lemurs, lorises and galagos. A toothcomb consists of long, flat forward-angled teeth, and includes the lower incisors and the canine teeth...

.

Distribution

The distribution of the Bornean slow loris is centered on the island of Borneo
Borneo
Borneo is the third largest island in the world and is located north of Java Island, Indonesia, at the geographic centre of Maritime Southeast Asia....

, where it occurs in Brunei
Brunei
Brunei , officially the State of Brunei Darussalam or the Nation of Brunei, the Abode of Peace , is a sovereign state located on the north coast of the island of Borneo, in Southeast Asia...

, Kalimantan
Kalimantan
In English, the term Kalimantan refers to the Indonesian portion of the island of Borneo, while in Indonesian, the term "Kalimantan" refers to the whole island of Borneo....

 (the Indonesian part of Borneo), and Sabah
Sabah
Sabah is one of 13 member states of Malaysia. It is located on the northern portion of the island of Borneo. It is the second largest state in the country after Sarawak, which it borders on its southwest. It also shares a border with the province of East Kalimantan of Indonesia in the south...

 and Sarawak
Sarawak
Sarawak is one of two Malaysian states on the island of Borneo. Known as Bumi Kenyalang , Sarawak is situated on the north-west of the island. It is the largest state in Malaysia followed by Sabah, the second largest state located to the North- East.The administrative capital is Kuching, which...

 (the Malaysian part of the island). The species is also found on the Indonesian islands of Bangka
Bangka Island
Bangka is an island lying east of Sumatra, Indonesia. Population 626,955. Area: c.4,600 sq mi .There is an additional small island named Pulau Bangka in northern Sulawesi, Indonesia.-Geography:...

 and Belitung
Belitung
Belitung, , is an island on the east coast of Sumatra, Indonesia in the Java Sea. The island is known for its pepper and for its tin. It was in the possession of the British from 1812 until the British ceded control of the island to the Dutch in the Anglo-Dutch Treaty of 1824...

, to the southwest of Borneo, in the Sulu Archipelago
Sulu Archipelago
The Sulu Archipelago is a chain of islands in the southwestern Philippines. This archipelago is considered to be part of the Moroland by the local rebel independence movement. This island group forms the northern limit of the Celebes Sea....

 of the southern Philippines
Philippines
The Philippines , officially known as the Republic of the Philippines , is a country in Southeast Asia in the western Pacific Ocean. To its north across the Luzon Strait lies Taiwan. West across the South China Sea sits Vietnam...

, and on the island of Banggi off Sabah. In the Sulu Archipelago, it occurs in the Tawi-Tawi Group, in the west of the archipelago, including the islands of Tawi-Tawi
Tawi-Tawi
Tawi-Tawi is an island province of the Philippines located in the Autonomous Region in Muslim Mindanao . The capital of Tawi-Tawi is Bongao. The province is the southernmost of the country sharing sea borders with the Malaysian State of Sabah and the Indonesian East Kalimantan province. To the...

, Bongao, Sanga-Sanga, Simunul, and possibly other small islands. It does not occur on the island of Jolo
Jolo
Jolo may refer to:* Jolo Island* Jolo, Sulu* Jolo, West Virginia* Jolo is also the nickname of Swedish author Jan Olof Olsson....

, further to the east, and although it has been recorded from Mindanao
Mindanao
Mindanao is the second largest and easternmost island in the Philippines. It is also the name of one of the three island groups in the country, which consists of the island of Mindanao and smaller surrounding islands. The other two are Luzon and the Visayas. The island of Mindanao is called The...

—a large island in the southern Philippines—the record was in error. The species may be extinct on some Philippine islands, but is likely to persist on the smaller islands. Because the species is so popular as a pet, zoologists Guy Musser
Guy Musser
Guy Graham Musser is an American zoologist. His main research field is the subfamily Murinae, in which he has described many new species.Musser was born in Salt Lake City, Utah...

 and Lawrence Heaney suggested in 1985 that the Philippine populations may have been introduced
Introduced species
An introduced species — or neozoon, alien, exotic, non-indigenous, or non-native species, or simply an introduction, is a species living outside its indigenous or native distributional range, and has arrived in an ecosystem or plant community by human activity, either deliberate or accidental...

 there by humans.

Fossils of this species have been found in the Late Pleistocene
Late Pleistocene
The Late Pleistocene is a stage of the Pleistocene Epoch. The beginning of the stage is defined by the base of the Eemian interglacial phase before the final glacial episode of the Pleistocene 126,000 ± 5,000 years ago. The end of the stage is defined exactly at 10,000 Carbon-14 years BP...

 site of Niah in Sarawak
Sarawak
Sarawak is one of two Malaysian states on the island of Borneo. Known as Bumi Kenyalang , Sarawak is situated on the north-west of the island. It is the largest state in Malaysia followed by Sabah, the second largest state located to the North- East.The administrative capital is Kuching, which...

.

Habitat and ecology

The Bornean slow loris is the least studied of Indonesia's slow lorises. In a field study at the Sabangau National Park
Sabangau National Park
Sabangau National Park is a national park in Central Kalimantan, a province of Indonesia in Kalimantan, the Indonesian part of the island of Borneo.- Geography :...

 in Central Kalimantan
Central Kalimantan
Central Kalimantan is a province of Indonesia, one of four in Kalimantan - the Indonesian part of the island of Borneo. Its provincial capital is Palangka Raya.The province has a population of just over 2.2 million at the 2010 Census...

, only 12 slow loris sightings were made over a 75-day period. All were seen in the trees at heights of 15 –. They were encountered singly, as mother and offspring, or in adult trios. Of the two trios, both were on fruiting trees, Calophyllum hosei and Syzygium
Syzygium
Syzygium is a genus of flowering plants that belongs to the myrtle family, Myrtaceae. The genus comprises about 1100 species, and has a native range that extends from Africa and Madagascar through southern Asia east through the Pacific...

cf. nigricans. In another survey conducted at Wehea, East Kalimantan
East Kalimantan
East Kalimantan is the second largest Indonesian province, located on the Kalimantan region on the east of Borneo island. The resource-rich province has two major cities, Samarinda and Balikpapan...

, only one Bornean slow loris—seen at a height of 30 m (98.4 ft)—was encountered in an area of more than 30 km2. Other surveys confirm that the animal is difficult to locate, and occurs in low densities
Population density
Population density is a measurement of population per unit area or unit volume. It is frequently applied to living organisms, and particularly to humans...

.

The species occurs in primary and secondary lowland forest, gardens, and plantations, at elevations between 35 –. Interviews conducted with Philippines locals indicate that it is commonly seen in citrus trees (calamansi), and may be tolerant of a variety of habitats. It is nocturnal, and almost entirely arboreal. Although data on diet is limited, based on cranial size and morphology
Morphology (biology)
In biology, morphology is a branch of bioscience dealing with the study of the form and structure of organisms and their specific structural features....

, the Bornean slow loris is suspected to be one of the more insectivorous
Insectivore
An insectivore is a type of carnivore with a diet that consists chiefly of insects and similar small creatures. An alternate term is entomophage, which also refers to the human practice of eating insects....

 slow loris species. It has also been observed feeding on the gum
Gum (botany)
Gum is a sap or other resinous material associated with certain species of the plant kingdom. This material is often polysaccharide-based and most frequently is associated with woody plants, particularly under the bark or as a seed coating...

 from an unidentified liana (a long-stemmed woody vine).

Conservation

In a 2005 report on the effect of logging on wildlife conservation in Indonesia, the authors claimed the Bornean slow loris to be "common" throughout Borneo. However, as pointed out by Nekaris and colleagues, this assessment was based on field research data and historic museum specimens, and cannot be considered reliable, as "loris ‘presence’ is usually not determined first-hand, and it also cannot be presumed that lorises still occur in areas from where they were once collected." The species appears to be uncommon throughout its range, including a very limited distribution in the Philippines. Surveys have demonstrated that, compared to other slow loris species, the Bornean slow loris is rare, and sparsely distributed throughout its range.

The Bornean slow loris is listed in CITES Appendix I, which prevents international commercial trade; it is also protected by Indonesian law. The species is often confused with other slow lorises in animal rescue centers, as it is not well-covered in field guides. The species occurs in a number of protected area
Protected area
Protected areas are locations which receive protection because of their recognised natural, ecological and/or cultural values. There are several kinds of protected areas, which vary by level of protection depending on the enabling laws of each country or the regulations of the international...

s throughout its range, including some fragmented forests. Threats to the species include the illegal local exotic pet
Exotic pet
An exotic pet is a rare or unusual animal pet, or an animal kept as a pet which is not commonly thought of as a pet.The definition is an evolving one; some rodents, reptiles, and amphibians have become firmly enough established in the world of animal fancy to no longer be considered exotic...

 trade and habitat loss due to burning and conversion to palm oil plantations. Additionally, uncontrolled release of pets in some areas is also a threat to the species.

The Bornean slow loris is among the least threatened of the slow lorises, and its situation is considered to be good due to its presence in a high percentage of "low risk" areas on Borneo. It is classified as "Vulnerable
Vulnerable species
On 30 January 2010, the IUCN Red List of Threatened Species identified 9694 Vulnerable species, subspecies and varieties, stocks and sub-populations.-References:...

" by the IUCN, who consider there to have been a greater than 30% reduction in population over the past 21–24 years, based on harvesting for the pet trade and extensive habitat loss. Using combined data from studies conducted in Danum, Malaysia, and Sabangau, the encounter rate was 0.12 individuals/kilometer.

Literature cited

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