The book of earth: lemore ( Lorisidae )

The Bengal slow loris (Nycticebus bengalensis) 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 (10 to 15 in) from head to tail and weighing between 1 and 2.1 kg (2.2 and 4.6 lb). 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 (a scent gland in its arm) differs chemically from that of other slow loris species and may be used to communicate information about sex, age, health, and social status.

The Bengal slow loris is nocturnal and arboreal, occurring in both evergreen and deciduous forests. It prefers rainforests with dense canopies, and its presence in its native habitat indicates a healthy ecosystem. It is a seed disperser and pollinator, as well as a prey item for carnivores. Its diet primarily consists of fruit, but also includes insects, tree gum, snails, and small vertebrates. In winter, it relies on plant exudates, such as sap and tree gum. The species lives in small family groups, marks its territory with urine, and sleeps during the day by curling up in dense vegetation or in tree holes. It is a seasonal breeder, reproducing once every 12–18 months and usually giving birth to a single offspring. For the first three months, mothers carry their offspring, which reach sexual maturity at around 20 months. The Bengal slow loris can live up to 20 years.

The species is listed as "Vulnerable" on the IUCN Red List, and is threatened with extinction due to a growing demand in the exotic pet trade and traditional medicine. It is one of the most common animals sold in local animal markets. In traditional medicine, it is primarily used by wealthy to middle-class, urban women following childbirth, but also to treat stomach problems, broken bones, and sexually transmitted diseases. It is also hunted for food and suffers from habitat loss. Wild populations have declined severely, and it is locally extinct in several regions. It is found within many protected areas throughout its range, but this does not protect them from rampant poaching and illegal logging. Critical conservation issues for this species include enhancing protection measures, stricter enforcement of wildlife protection laws, and increased connectivity between fragmented protected areas.

Taxonomy and phylogeny

An old drawing of a Bengal slow loris sitting on a branch with another moving between branches in the background
The Bengal slow loris was long considered a subspecies until it was recognized as a distinct species in 2001.
Nycticebus bengalensis, commonly known as the Bengal slow loris or northern slow loris, is a strepsirrhine primate in the slow loris genus, Nycticebus. Formerly considered a subspecies of the Sunda slow loris (N. coucang), it was recognized as a distinct species in 2001 by taxonomist and primatologist Colin Groves. It is difficult to distinguish from the other species in its genus.

To help clarify species and subspecies boundaries, and to establish whether morphology-based classifications were consistent with evolutionary relationships, the phylogenetic relationships within the genus Nycticebus have been investigated using DNA sequences derived from the mitochondrial markers D loop and cytochrome b. Although most of the recognized lineages of Nycticebus (including N. pygmaeus, N. menagensis and N. javanicus) were shown to be genetically distinct—the analysis suggested that DNA sequences from some individuals of N. coucang and N. bengalensis apparently share a closer evolutionary relationship with each other than with members of their own species. The authors suggest that this result may be explained by introgressive hybridization, as the tested individuals of these two taxa originated from a region of sympatry in southern Thailand; the precise origin of one of the N. coucang individuals was not known. This hypothesis was corroborated by a 2007 study that compared the variations in mitochondrial DNA sequences between N. bengalensis and N. coucang, and suggested that there has been gene flow between the two species.

Anatomy and physiology

The Bengal slow loris has large eyes, round head, and short ears, as shown in this illustration from the mid-19th century.
The Bengal slow loris is the largest species of slow loris, weighing 1 to 2.1 kg (2.2 to 4.6 lb), and measuring between 26 and 38 cm (10 and 15 in) from head to tail. It has a skull length of more than 62 mm (2.4 in). It has dense, woolly, brown-gray fur on its back and white fur on its underside. It also has a clear dark stripe that runs up to the top of its head, but does not extend laterally towards the ears. Its forearm and hand are almost white. The limbs of the pelvis vary in color from brown to nearly white, and the feet are always pale. Moulting may cause seasonal variations in the color of the dorsal surface. Like other slow lorises, its tail is vestigial and it has a 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. Its eyes reflect a bright orange eye shine. 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" that the animal uses for scratching and grooming, while the other nails are straight.

In addition to being smaller than the Bengal slow loris, the sympatric Sunda slow loris also differs in its coloring: it does not have the pale areas of the head, nape and shoulders, and its overall color is a tawny- or golden-brown. The pygmy slow loris (N. pygmaeus) is much smaller, with a skull length less than 55 mm (2.2 in). It also lacks the dark dorsal stripe of the Bengal slow loris, has dark brown fur, and longer ears.

The Bengal slow loris 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. The oil has been analyzed using gas chromatography coupled to mass spectrometry, and it has been shown that almost half of the several dozen volatile or semi-volatile chemicals present do not occur in the closely related pygmy slow loris. The most predominant component was the phenolic compound m-cresol. The authors of the study suggest that the chemically complex oils may help the lorises communicate with each other, allowing them to transmit by scent information about sex, age, health and nutritional status, and dominance.

Behavior and ecology

The Bengal slow loris will gouge the bark of the bastard myrobala tree to obtain exudates, an important food source—especially in winter seasons.
The preferred habitats of the Bengal slow loris range across tropical and subtropical regions, and include evergreen and semi-evergreen rainforests with forest edges and continuous, dense canopies. It can also be found in bamboo groves. It prefers habitats with larger diameter, tall trees with a large crown depth (defined as the length along the main axis from the tree tip to the base of the crown); these areas are typically associated with greater food abundance, and decreased risk of predation. Because of its preference for dense forests, it acts as a good indicator of the ecosystem's health.

The species acts as an important seed disperser and pollinator, as well as a prey item for several carnivores. The Bengal slow loris feeds on plant exudates such as sap, gums, resins, and latexes, particularly those from the family Fabaceae. Even though the species does not have keeled nails, it will scrape the plant, actively breaking its surface; this behavior resembles that of marmosets and the fork-marked lemurs. Exudates are also obtained by gouging holes in the bark. The winter food supply consists almost entirely of plant exudates. The bastard myrobala (Terminalia belerica), a deciduous tree common in Southeast Asia, is a preferred source for exudates, but it has also been observed taking plant exudates from a number of families: Moraceae (Artocarpus), Magnoliaceae (Manglietia), Fabaceae (Acacia, Bauhinia), Lecythidaceae (Careya arborea), and Sterculiaceae (Pterospermum). Although it will feed on large insects (such as katydids and crickets), gum, snails, small birds, and reptiles, it is primarily frugivorous. Lianas of the flowering plant genus Bauhinia are a commonly used food source.

A nocturnal animal, the Bengal slow loris has excellent night vision, enhanced by a tapetum lucidum—a layer of tissue in the eye that reflects visible light back through the retina. It sleeps during the day curled up in a ball in dense vegetation or in tree holes. Males and females mark their territory with urine. The species is known to live in small family groups. Animals may practice social grooming.

The Bengal slow loris is not a seasonal breeder, unlike the pygmy slow loris. Females in an estrous cycle attract males with a loud whistle. Females reproduce every 12–18 months and have a six-month gestation. Because they are not seasonal breeders, females could become pregnant when their offspring are approximately 6 months old, making possible for females to produce two offspring per year. Females typically give birth to a single offspring, although twins rarely occur. This differs from the sympatric pygmy slow loris, which commonly has twins. The mother carries her young about three months before they become independent, although they may be temporarily left on branches while the mother searches for food. Sexual maturity is reached at approximately 20 months of age. The species is known to live up to 20 years.

Distribution

The species has the largest geographic range of all slow loris species and is native to Northeast India, Bangladesh and Indochina (Cambodia, Laos, Burma, Vietnam, southern China and Thailand). It is the only nocturnal primate found in the northeast Indian states, which include Assam, Arunachal Pradesh, Mizoram, Nagaland, Meghalaya, Manipur and Tripura. It is found in parts of Yunnan and in southwest Guangxi in China, and has been recorded in the Chittagong Hill Tracts in Bangladesh. It is known from 24 protected areas in Vietnam, and is distributed across most of Thailand. In Burma, it has been reported from Bhamo, Sumprabum, Kindat, Chin Hills, Pathein, Thaungdaung and Pegu; populations in Laos have been recorded in the north, central, and southern portions of the country.

The Bengal slow loris is sympatric (shares its range) with the pygmy slow loris in southeast of China, Vietnam, and Laos. The Bengal slow loris is also sympatric with the Sunda slow loris on the southern peninsula of Thailand. In 2001, Groves reported the existence of hybrids between these two species in this region.

Conservation

See also: Conservation of slow lorises
Listed as "Data Deficient" as recently as 2006 on the IUCN Red List, The Bengal slow loris was evaluated in 2008 by the International Union for Conservation of Nature (IUCN) as "Vulnerable"—a decision based solely on habitat loss due to lack of sufficient field data. It is found within numerous protected areas within its range; however, poaching and illegal logging are rampant while conservation measures are not species-specific. The species can be found in at least 43 protected areas in Northeast India, 14 conservation areas in Laos, and 24 protected areas in Vietnam. It can be found at Lawachara National Park in Bangladesh, and 80% of its range in China is protected. The species has been listed in Schedule I of the Indian Wildlife Protection Act of 1972, and in June 2007, it was transferred along with all other slow loris species to CITES Appendix I, which forbids international commercial trade.

The most severe threats facing the species are the wildlife trade (trapping for exotic pets and use in traditional medicine) and deforestation. Slash and burn agriculture has also resulted in the destruction of its habitat, and road construction is another factor in its decline. Hunting has been found to be most severe when nearby urban human populations increase. Enhancing protection measures, enforcing current wildlife protection laws, and improving the connectivity between protected areas are factors considered critical to ensure the survival of this species.

The species is commonly sold as a pet and to zoos throughout Southeast Asia. In Cambodia, it was reported in 2006 as one of the most common mammals found in shops and stalls, found in the hundreds and selling for US$0.85 to US$6.25. In the same year, it was found selling for US$2.50 to US$6.30 at bazaars in China (Mengla County in Yunnan Province) and US$70 in Thailand. The Bengal slow loris is used in traditional medicine in all of these countries, selling for US$15 in Vietnam, and is also eaten in Vietnam. The animal is predominantly used to prepare treatments for women after childbirth, stomach problems, healing wounds and broken bones, and in the treatment of sexually transmitted diseases. Primary users are wealthy to middle-class women in urban areas.

Throughout its geographic range, slow lorises are in serious decline. Their habitat has been seriously degraded, and growing human populations will add increasing pressure. In countries like Bangladesh, only 9% of the original forest cover was still present in 2000. In northeastern Cambodia, forests are being cleared at an increasing rate, with a loss of 6% of the natural forest between 1999 and 2000. Within those same years, Myanmar and Thailand lost 14% and 26% of their natural forest, respectively. In Vietnam, only 30% of the original forest cover remains due to the deforestation caused by the Vietnam War, and only 10% of that includes closed-canopy forests. Habitat destruction remains rampant, and all slow loris populations within its borders are significantly depleted. Populations have been declared locally extinct in southern Quảng Nam Province and parts of the highlands, and the same is expected in Song Thanh and Kon Cha Rang nature reserves.

In India, dense forest canopy has been depleted by as much as 55% in some areas and is rapidly disappearing. As early as 1987, the Indo-China region had reportedly lost 75% of the natural habitat for slow lorises. In 1992, the population size was estimated between 16,000 and 17,000 individuals, based on available habitat; however, recent publications report that few individuals remain due a reduced geographic range. The Bengal slow loris may be restricted to a few isolated populations and is in serious threat of becoming locally extinct in parts of Assam and Meghalaya. In Arunachal Pradesh, its population is declining and under threat.

Population density has been estimated between 0.03 and 0.33 individuals per km2 in Assam, India according to a study published in 2006. A survey in 2007 at the Thrisna Wildlife Sanctuary and Sipahijola Wildlife Sanctuary in Tripura, India yielded an encounter rate of 0.22 individuals/km, with seven of nine sightings occurring within 1.71 km2 (0.66 sq mi) and most of the animals found at a height of 8–15 m (26–49 ft) and near the interior of wet, deciduous forest. In 2008, the species abundance was measured at 0.18 individuals/km at Gibbon Wildlife Sanctuary in Assam.

Since the 1990s, China's forests have declined significantly. In Yunnan and Guangxi provinces, primary forests are few and isolated, and secondary forests have been severely degraded. Yunnan has lost 42% of its forests and 2,000 or less slow lorises remain. In Guangxi, the Bengal slow loris is nearly extinct; it has been extirpated in Ningming County and only a few individuals are left in Jingxi, Longzhou and Pingxiang.

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The Javan slow loris (Nycticebus javanicus) 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.

Its forehead has a prominent white diamond pattern, which consists of a distinct stripe that runs over its head and forks towards the eyes and ears. The Javan slow loris weighs between 565 and 687 g (1.25 and 1.51 lb) and has a head-body length of about 293 mm (11.5 in). Like all lorises it is arboreal, and moves slowly across vines and lianas instead of jumping from tree to tree. Its habitat includes primary and secondary forests, but it can also be found in bamboo and mangrove forests, and on chocolate plantations. Its diet typically consists of fruit, tree gum, lizards and eggs. It sleeps on exposed branches, sometimes in groups, and is usually seen alone or in pairs.

The Javan slow loris population is in sharp decline because of poaching for the exotic pet trade, and sometimes for traditional medicine. Remaining populations have low densities, and habitat loss is a major threat. For these reasons the International Union for Conservation of Nature (IUCN) lists its status as critically endangered, and it has also been included on the 2008–2010 list of "The World's 25 Most Endangered Primates". It is protected by Indonesian law and, since June 2007, is listed under CITES Appendix I. Despite these protections, as well as its presence in several protected areas, poaching continues; the wildlife protection laws are rarely enforced at the local level.

Taxonomy and phylogeny

The Javan slow loris (Nycticebus javanicus) was first described scientifically in 1812, by the French naturalist Étienne Geoffroy Saint-Hilaire. The species name javanicus refers to its place of origin. However, the species was not recognized for long; by 1840, René Primevère Lesson classified it as one of several varieties of a single species of slow loris, which he called Bradylemur tardigradus. In 1921, Oldfield Thomas named a second species of slow loris from Java, Nycticebus ornatus.

In his 1971 review of slow loris taxonomy, taxonomist and primatologist Colin Groves recognized the Javan slow loris as a subspecies, Nycticebus coucang javanicus, of the Sunda slow loris (N. coucang), with ornatus as a synonym. It was first recognized as a distinct species again in a 2000 Indonesian field guide on primates by Jatna Supriatna and Edy Hendras Wahyono. In 2008, Groves and Ibnu Maryanto promoted it to species status, based on an analysis of cranial morphology and characteristics of pelage. Molecular analysis of DNA sequences of the D-loop and cytochrome b genes demonstrated it to be genetically distinct from other slow loris species; phylogenetically, it is sister to a clade containing the Bengal slow loris (N. bengalensis) and the Sunda slow loris. Due to its close resemblance to neighboring slow loris species, even rescue centers have been known to misidentify it.

There are two forms of the Javan slow loris, distinguished mainly by differences in hair length. These have occasionally been recognized as separate species, N. javanicus and N. ornatus, but are currently both classified as a single species, although their exact taxonomic status remains unclear.

Anatomy and physiology

The Javan slow loris weighs between 565 and 687 g (1.25 and 1.51 lb) and is similar in appearance to the largest slow loris, the Bengal slow loris. Its face and back are marked with a distinct stripe that runs over the crown and forks, leading to the eyes and ears, which leaves a white diamond pattern on the forehead. Its color is yellowish-gray. In contrast, its head, neck, and shoulders have cream hues. Like the Bornean slow loris (N. menagensis), it lacks the second incisor (I2) in its dentition.

A Javan slow loris clings perpendicularly to a vertical strand of bamboo.
The Javan slow loris has a distinct stripe that runs the length of its back and forks at the crown, leading to the eyes and ears.
The Javan slow loris is larger than both of the other Indonesian slow lorises, the Sunda slow loris and the Bornean slow loris. Based on averages determined from six specimens obtained from the illegal wildlife trade in Java, other morphometric parameters are as follows: head length, 59.2 mm; muzzle length, 19.9 mm; head breadth, 43.6 mm; body breadth, 250.8 mm; head and body length, 293.1 mm; chest girth, 190.8 mm; dark percentage girth (girth measurement of zone with dark dorsal hair, measured as a percent of girth circumference), 48.0 mm; neck circumference, 136.7 mm; tail length, 20.4 mm; humerus length, 67.2 mm; radius length, 71.8 mm; femur length, 83.2 mm; tibia length, 85.9 mm; hand span, 59.1 mm; foot span, 70.3 mm; and ear length, 16.8 mm.

The ornatus morphotype is most reliably distinguished by its longer fur, averaging 26.8 mm compared to 22.4 mm in javanicus. Other distinguishing characteristics include overall color (generally light brown in ornatus compared with brown to reddish in javanicus), and amount of brown coloring in the fur (ornatus has less brown than javanicus, resulting in a lighter-colored ventral region).

In the 1860s, the brain of the Javan slow loris was examined by William Henry Flower, a comparative anatomist who specialized in the primate brain. In addition to detailing the organization, shape, and measurements of its brain, he noted that the form and surface markings were comparable to that of lemurs. He argued against grouping strepsirrhines with Insectivora (a now-abandoned biological grouping) and noted that the brain had features transitional between other primates and "inferior" mammals such as bats and carnivorans.

Behavior and ecology

Like other lorises, the Javan slow loris is nocturnal and arboreal, relying on vines and lianas. However, the animal has been observed moving on the ground to cross open spaces in disturbed habitat. It moves through the canopy at heights between 3 and 22 m (9.8 and 72.2 ft) and is often encountered at heights between 1.5 and 9.5 m (4.9 and 31.2 ft).

The Javan slow loris will eat fruit, lizards, eggs, and chocolate seeds. It is also known to eat the gum of trees of the genus Albizia, in the legume family, Fabaceae, as well as from the palm genus Arenga (family Arecaceae). Javan slow lorises are seen alone or in pairs and are sometimes found sleeping in groups. Instead of sleeping in nest holes, they sleep curled up on branches. Like other slow lorises, the Javan slow loris has a distinctive call that resembles a high-frequency whistle. The species is a host for the parasitic flatworm, Phaneropsolus oviforme.

Distribution

The species is found only on the western and central portion of the island of Java in Indonesia. Its presence has been confirmed in the Dieng Mountains, and it is known to be found in low densities at Gunung Gede Pangrango National Park (in montane cloud forests) and Mount Halimun Salak National Park, often only where human disturbance is minimal. It inhabits both primary and secondary disturbed forest, and can be found from sea level to 1,600 m (5,200 ft), although it is more commonly found at higher elevations since lower elevations tend to be deforested. A study in 2000 showed that in addition to primary and secondary forest, the Javan slow loris could be found in bamboo forests, mangrove forests, and on plantations—particularly chocolate plantations. In 2008, they were observed in West Java to occupy mixed-crop home gardens, tolerating high levels of human disturbance.

Conservation

See also: Conservation of slow lorises
Three Javan slow lorises sit curled up on the bottom of a wired cage
The Javan slow loris is commonly sold as a pet in the markets of Indonesia, despite local laws and CITES Appendix I protection.
The Javan slow loris is listed by the International Union for Conservation of Nature (IUCN) as "critically endangered," primarily due to a rapid decline in population. For the 21–24 years prior to its 2008 assessment by the IUCN—which corresponds to three generations for the species—its numbers had dropped by at least 50%. Population data for the species is sparse, but a few studies have shown a low population density of 0.20 to 0.02 individuals per km2.

Its numbers are still decreasing, primarily because of poaching. In Indonesia, it is sometimes used in traditional medicine, because of myths of it having magical and curative properties, but it is more frequently sold as an exotic pet. The species is easily captured because of its slow movement, nocturnal habits, and its tendency to sleep on exposed branches. They are both actively sought for the pet trade and collected opportunistically when felling forests. Its habitat is also in decline, although most of the habitat loss occurred by the mid-1980s. Within its range, human land use is intense. Environmental niche modelling indicates that the Javan slow loris is more threatened by habitat loss than other slow loris species. For these reasons, the Javan slow loris has been included on "The World's 25 Most Endangered Primates" published by the IUCN Species Survival Commission Primate Specialist Group (IUCN/SSC PSG), the International Primatological Society (IPS), and Conservation International (CI).

Along with all other slow lorises, the Javan slow loris was elevated from CITES Appendix II to CITES Appendix I in June 2007, offering it increased protection from commercial trade. It is also protected by Indonesian law, but according to loris researchers Nekaris and Jaffe, "effective law enforcement with respect to wildlife protection laws is all but non-existent in Indonesia". The species can be found in several protected areas, but its numbers are uncertain. Captive collections of the Javan slow loris can be found in Prague, Czech Republic, Jakarta, Indonesia, and Singapore.

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"Bornean slow loris" redirects here. For other slow lorises of Borneo that have shared that name, see Bornean slow loris (disambiguation).
The Bornean slow loris (Nycticebus borneanus) 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

Female, from Nanga Tayap, Ketapang Regency
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.

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This page is about a species that has been called Bornean slow loris. For the present-day Bornean slow loris, see Nycticebus borneanus. For other slow lorises of Borneo, see Bornean slow loris (disambiguation).
The Philippine slow loris (Nycticebus menagensis) 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.

Weighing 265–300 grams (9.3–10.6 oz), 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. Like other slow lorises, it has a vestigial tail, round head, short ears, a curved grooming claw for grooming, and a gland that produces an oily toxin that the animal uses for defense. The Philippine slow loris is arboreal, nocturnal, and occurs in low densities, making it difficult to locate. It is also the least studied of Indonesia's slow lorises. It is found at elevations between 35–100 meters (115–328 ft) in primary and secondary lowland forest, gardens, and plantations. Information about its diet is limited, but it is suspected to be one of the more insectivorous slow loris species, and is also known to eat gum from woody plants.

The Bornean slow loris species complex – including the Philippine slow loris – was classified as "Vulnerable" by the International Union for Conservation of Nature (IUCN) in 2008, is included in CITES Appendix I, which prevents international commercial trade, and is protected by Indonesian law. Prior to being divided into four distinct species in 2013, it was found in numerous protected areas within its range, making it the least threatened of the slow lorises. However, since the taxonomic split, it may face a higher risk of extinction. It is sparsely distributed throughout its range and is threatened by illegal wildlife trade, including the exotic pet 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 near Tataan, Tawi-Tawi Island, in the Philippines, however this type specimen is missing as of 2013. 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. 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:

I now come to the curious mammal of which I enclose description. Shortly before we left for Tawi Tawi the Jesuit priest here, Padre Marche, informed us that just before our arrival he had made a trip to Tawi Tawi, and had bought of the Moros there a curious animal. He said it has the face of a bear, the hands of a monkey, moved like a sloth, and was called "cocam" by the natives ... I believe nothing of this kind has been found in the Philippines before, and it makes an important addition to the rather meager list of Philippine mammals. It is evidently one of the Lemuridae, but as generic characteristics are not given in the book I have, I cannot go farther.

— Dean C. Worcester
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 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 1953 book Primates: Comparative Anatomy and Taxonomy, the primatologist William Charles Osman Hill placed all the slow lorises in one species, N. coucang. In 1971, however, Colin Groves recognized the pygmy slow loris (N. pygmaeus) as a separate species, and divided N. coucang into four subspecies, including Nycticebus coucang menagensis. Nycticebus menagensis was elevated to the species level in 2006 when molecular analysis of DNA sequences of the D-loop and the cytochrome b gene demonstrated it to be genetically distinct from N. coucang. The genetic evidence was corroborated by both a previous study (1998) on morphology (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.

Early in its own taxonomic history, distinguishing coloration patterns and size differences resulted in the division of Nycticebus menagensis into four subspecies: N. m. bancanus, N. m. borneanus, N. m. menagensis, and N. m. philippinus. These later became taxonomic synonyms, although in 2013 N. bancanus and N. borneanus were elevated to species status based on unique facial markings. Furthermore, a new species—N. kayan—was also identified within the Bornean population. The southern Philippine slow lorises (N. m. philippinus or N. philippinus) identified by Spanish zoologist Ángel Cabrera in 1908 remains a synonym of N. menagensis, since the 2013 study was unable to find its type specimen and found no distinguishable characteristics between the two populations. The syntype for N. menagensis was collected by Lydekker in 1893 from the Philippine island of Tawi-Tawi, giving menagensis precedence over philippinus as the name of the species, under the Principle of Priority.

Physical description

Like all slow lorises, the tail of the Philippine slow loris is vestigial and it has a 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. 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 the animal 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. 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.

The body weight of this species is typically in the range of 265–325 grams (9.3–11.5 oz), although weights of up to 700 grams (25 oz) have been recorded. The body length averages 274.2 mm (10.80 in),, and its skull length ranges between 54.5 and 56.5 mm (2.15 and 2.22 in), roughly intermediate in size between the smaller pygmy slow loris and the larger Sunda slow loris. The slow lorises of Borneo are among the smallest of its genus, but this species can be distinguished from the others by its pale golden to red fur, low-contrast markings on its face and head, and the consistent lack of a second upper incisor. The rings around the eyes are either rounded or diffused-edged on top, while the bottom occasionally extends down below the zygomatic arch. The stripe between its eyes is narrow, the ears usually lack fur, the patch on the top of the head is mostly diffused, and the band of fur in front of the ears varies in width. In comparison to the other three species of slow lorises on Borneo, both N. menagensis and N. kayan have a pale body coloration, but this species has pale, very lightly-contrasting facial markings, with markedly less contrast than the dark, high-contrast face mask of N. kayan. Additionally, this species has short, unfluffed body hair, in contrast to the longer, fluffier body hair of N. kayan.

Distribution

N. menagensis is found primarily in coastal and lowland areas in northern and eastern Borneo, in the provinces of Brunei, Sabah, and East Kalimantan. It is also found on the southern Philippine Islands, known as the Sulu Archipelago, and may be found on other nearby islands, such as the Banggi Island 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, Bongao, Sanga-Sanga, Simunul, and possibly other small islands. It does not occur on the island of Jolo or further to the east, as was claimed by one erroneous report from Mindanao. 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 and Lawrence Heaney suggested in 1985 that the Philippine populations may have been introduced there by humans.

Of the four slow lorises living on Borneo, N. menagensis is believed to be partly sympatric with N. kayan only, with ranges that overlap in East Kalimantan and Sabah. Fossils of this species have been found in the Late Pleistocene site of Niah in Sarawak.

Habitat and ecology

The slow lorises of Borneo are the least studied of Indonesia's slow lorises. In a field study at the Sabangau National Park in Central Kalimantan, only 12 slow loris sightings were made over a 75-day period. All were seen in the trees at heights of 15–20 m (49–66 ft). 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 cf. nigricans. In another survey conducted at Wehea Forest, East Kalimantan, only one Philippine slow loris—seen at a height of 30 m (98 ft)—was encountered in an area of more than 30 km2 (12 sq mi). Other surveys confirm that the animal is difficult to locate, and occurs in low densities.

The species occurs in primary and secondary lowland forest, gardens, and plantations, at elevations between 35–100 m (115–328 ft). 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, the Philippine slow loris is suspected to be one of the more insectivorous slow loris species. It has also been observed feeding on the gum from an unidentified liana (a long-stemmed woody vine).

Conservation

See also: Conservation of slow lorises
In a 2005 report on the effect of logging on wildlife conservation in Indonesia, the authors claimed N. menagensis 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 Philippine slow loris is rare, and sparsely distributed throughout its range.

N. menagensis 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 areas throughout its range, including some fragmented forests. Threats to the species include the illegal local exotic pet 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.

Prior to being split into four species in 2013, the Bornean slow loris species complex was among the least threatened of the slow lorises, and its situation was considered to be good due to its presence in a high percentage of "low risk" areas on Borneo. It was classified as "Vulnerable" by the IUCN, who consider there to have been a greater than 30% reduction in population between roughly 1984 and 2008, based on harvesting for the pet trade and extensive habitat loss. Because that species complex has been divided into four distinct species since the 2008 IUCN assessment, each of the new species likely 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.

source - Wikipedia

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