Slow loris

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Slow lorises[1]
Sunda Slow Loris
Nycticebus coucang
CITES Appendix I (CITES)[2]
Scientific classification
Kingdom:
Animalia
Phylum:
Chordata
Class:
Mammalia
Order:
Primates
Family:
Lorisidae
Subfamily:
Lorinae
Genus:
Nycticebus

Type species
Tardigradus coucang
Boddaert, 1785
Species

Nycticebus coucang
Nycticebus bengalensis
Nycticebus pygmaeus
Nycticebus javanicus
Nycticebus menagensis
†? Nycticebus linglom

Range map showing ranges of the three species of slow loris: the Sunda Loris (N. coucang) in Thailand, Malaysia, and Indonesia; the Bengal Slow Loris (N. bengalensis) in east India, China, Bangladesh, Bhutan, Burma, Thailand, Laos, Vietnam, and Cambodia; and the Pygmy Slow Loris (N. pygmaeus) in Vietnam and Laos.
Distribution of Nycticebus spp.
red = N. pygmaeus; blue = N. bengalensis;
brown = N. coucang, N. javanicus, & N. menagensis

Slow lorises (genus Nycticebus) are strepsirrhine primates found in South and Southeast Asia. They range from Northeast India to the southern Philippines and from the Yunnan province in China in the north to the island of Java in the south. There are currently five species recognized: Sunda slow loris (N. coucang), Bengal slow loris (N. bengalensis), pygmy slow loris (N. pygmaeus), Javan slow loris (N. javanicus), and the Bornean slow loris (N. menagensis). Slow lorises are most closely related to other lorisids, such as slender lorises, pottos, the false potto, and angwantibos. They are also closely related to the remaining lorisiforms—the various types of galago—as well as the lemurs of Madagascar. Their evolutionary history is unclear since their fossil record is patchy and molecular clock studies have given unclear or contradictory results.

Like other living ("crown") strepsirrhines, these nocturnal primates have a reflective layer in their eye called a tapetum lucidum, which enhances their night vision. They also have a rhinarium or "wet nose", which is a touch-based sense organ. Because of their close relation to lemurs and other lorisiforms, they also possess a toothcomb, which is used in personal and social grooming. Like nearly all prosimian ("pre-monkey") primates, they have a toilet-claw, which is also used in grooming. Slow lorises have a round head, narrow snout, large eyes, and distinctive coloration patterns. Their arms and legs are nearly equal in length, and their trunk is long, allowing them to twist and extend to nearby branches. The hands and feet of slow lorises have several adaptations that give them a pincer-like grip and enable them to grip branches for very long periods of time.

Slow lorises are the only primates to have a toxic bite. The toxin is produced by licking a gland on their arm, and the secretion mixes with its saliva to form the toxin. Little is known about their social structure, but they are known to communicate by scent-marking. Males are highly territorial. Slow lorises reproduce slowly, and the infants are initially parked on branches or carried by either parent. They are omnivores, eating small animals, fruit, tree gum, and other vegetation.

All slow loris species are listed as either "Vulnerable" or "Endangered" on the International Union for Conservation of Nature and Natural Resources Red List of Threatened Species (IUCN Red List) and are threatened by the wildlife trade and habitat loss. Although their habit is rapidly disappearing and becoming fragmented, making it nearly impossible for slow lorises to disperse between forest fragments, unsustainable demand from the exotic pet trade and traditional medicine has been the greatest cause for their decline. Despite local laws prohibiting the trade in slow lorises and slow loris products, as well as protection from international commercial trade under Appendix I, slow lorises are openly sold in animal markets in Southeast Asia and smuggled to countries like Japan, where they are popularized as pets in videos posted on YouTube. Slow lorises have their teeth cut or pulled out for the pet trade, and often die from infection, blood loss, poor handling, or poor nutrition.

Etymology

The genus Nycticebus, was named in 1812 by Étienne Geoffroy Saint-Hilaire[1] for its nocturnal behavior. The name derives from the Ancient Greek nycti- (νύξ, genitive form νυκτός), meaning "night" and cebus (κῆβος) meaning "monkey".[3]

The word "loris" was first used in 1765 by the French naturalist Georges-Louis Leclerc, Comte de Buffon as a close equivalent to the Dutch name, loeris. This etymology was later supported by the physician William Baird in the 1820s, who noted that the Dutch word loeris signified "a clown."[4]

Taxonomy and phylogeny

Slow lorises (genus Nycticebus) are strepsirrhine primates and are related to other living lorisiforms, such as slender lorises, pottos, false pottos, angwantibos, and galagos, as well as the lemurs of Madagascar.[5] The fossil record in both Asia and Africa is patchy, with most fossils dating back to the early Miocene, approximately 20 million years ago.[6] The evolutionary origins of crown strepsirrhines have traditionally been unclear due to a lack of sufficient fossil evidence, and often contradictory molecular clock, morphological, and biogeographic studies.[5] However in 2003, teeth and jaw fragments dating to the Late Middle Eocene (c. 40 million years ago) were found at the Birket Qarun Formation in the Egyptian Faiyum and were named Karanisia clarki. This fossil species is the oldest specimen to posses a toothcomb, a dental feature common among all living lorisiforms and lemurs, and has helped secure the evolutionary origins of the living (crown) strepsirrhines in Africa.[7] One of the simplest models of lorisiform evolution suggests that they diverged first from the lemurs and then from each other in Africa, with one group of lorisids later migrating to Asia and evolving into the slender and slow lorises of today.[8] Molecular clock analysis suggests that slow lorises may have started evolving into distinct species about 6 million years ago.[9]

Although Karanisia has been cautiously placed with angwantibos (lorisoids from Africa, genus Arctocebus) several other fossil species have been discovered in Asia that are more closely related to the slow lorises.[10] The fossil genus Nycticeboides dates from the late Miocene, and was found in Pakistan. Although it was comparable in size to the pygmy slow loris (Nycticebus pygmaeus), its teeth set it apart from living slow lorises.[11] In 1997, French paleontologists reported the discovery of a single tooth from a Miocene site in Thailand.[12] The tooth most closely resembled living slow lorises, and based on this limited material, it was named ? Nycticebus linglom, using open nomenclature (the preceding "?") to indicate the tentative nature of the assignment.[13]

The living slow lorises are generally considered to be most closely related to the slender lorises (genus Loris) of India, followed by the angwantibos, pottos, (genus Perodicticus), and the false potto (genus Pseudopotto), all from Central and West Africa.[10] However, the relationship between the African lorises and the Asian lorises is complicated by biogeography, strong similarities in morphology (with both slender and robust body forms existing in both Africa and Asia), and significant differences in genetics.[5]

Taxonomy

Phylogeny and relationships of N. coucang and related species based on mitochondrial DNA sequences.[14]

In his influential 1952 book Primates: Comparative Anatomy and Taxonomy, the primatologist William Charles Osman Hill placed all the slow lorises in one species, N. coucang.[15] In 1971 Colin Groves recognized the Pygmy Slow Loris (N. pygmaeus) as a separate species, and divided N. coucang into four subspecies,[16] while in 2001 Groves opined there were three species (N. coucang, N. pygmaeus, and N. bengalensis), and that N. coucang had three subspecies (Nycticebus coucang coucang, N. c. menagensis, and N. c. javanicus).[17] Species differentiation was based largely on differences in morphology, such as size, fur color, and head markings.[18]

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 by Chen and colleagues using DNA sequences derived from the mitochondrial markers D-loop and cytochrome b. Previous molecular analyses using karyotypes,[19] restriction enzymes,[20] and DNA sequences[21] were focused on understanding the relationships between species in the genus, not the phylogeny of the entire genus.[18] Although the sample size was limited, Chen and colleagues' 2006 publication showed that the lineages of Nycticebus can be generally divided into N. pygmaeus—which is deeply separated from the other species, and is thought to have been the first to diverge evolutionarily—and the other species. 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.[18] 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.[22]

Distribution and diversity

Slow lorises are found in South and Southeast Asia. Their collective range stretches from Northeast India through Indochina, east to the Sulu Archipelago (the small, southern islands of the Philippines), and south to the island of Java (including the Borneo, Sumatra, and many small nearby islands). They are found in India (Assam province), China (Yunnan province), Laos, Vietnam, Cambodia, Bangladesh, Burma, Thailand, Malaysia, the Philippines, and Indonesia. They have also been reported in Mindanao, the eastern-most island of the Philippines, although they were likely introduced there by humans.[23]

Slow lorises range across tropical and subtropical regions,[24] are found in primary and secondary rainforests, as well as bamboo groves, and mangrove forests.[25][26] They prefer forests with high, dense canopies,[27][24] although some species have also be found in disturbed habitats, such as chocolate plantations and mixed-crop home gardens.[25]

Physical description

Coloration patterns around the eyes differ between the slender lorises (middle two) and the slow lorises (top and bottom).

Slow lorises have a round head[28] because their skull is shorter from front to back compared to other prosimians.[29] Like other lorisids, its snout does not taper towards the front of the face as it does in lemurs. This makes the face less long and pointed.[30] Compared to the slender lorises, the snout of the slow loris is even less pointed.[28] A distinguishing feature of the slow loris skull is that the occipital bone is flattened and faces backward. The foramen magnum (hole through which the spinal cord enters) faces directly backward.[31] The brain of slow lorises has more folds (convolutions) than the brains of galagos.[32]

The ears are reduced in size,[5] sparsely covered in hair, and hidden in the fur.[27] Similar to the slender lorises, the fur around and directly above the eyes is dark. Unlike the slender lorises, however, the white stripe that separates the eye rings and broadens both on the tip of the nose and on the forehead while also fading out on the forehead.[27] Like other strepsirrhine primates, the nose and lip are covered by a moist skin called the rhinarium ("wet nose"), which is a sense organ.[33]

The eyes of slow lorises are forward-facing, which gives good depth-perception. Their eyes are large[34] and possess a tapetum lucidum, or reflective layer that improves low-light vision, but may also blur the images they see since the reflected light may interfere with the incoming light.[35] Slow lorises have monochromatic vision, meaning they see in shades of only one color. They lack the opsin gene that would allow them to detect short wavelength light, which includes the colors blue to green.[36]

The dental formula of slow lorises is 2.1.3.32.1.3.3 × 2 = 36, meaning that on each side of its mouth it has two upper and lower incisors, one upper and lower canine tooth, three upper and lower premolars, and three upper and lower molars, giving it a total of 36 permanent teeth.[37] Like all other crown strepsirrhines, their lower incisors and canine are procumbent (lie down and face outwards), forming a toothcomb, which is used for personal and social grooming.[37]

Slow lorises have a stout body,[27] and their tail is greatly reduced and hidden beneath the dense fur.[27][5] Their combined head and body length vary by species, with the smallest—the pygmy slow loris—measuring 18 to 21 cm (7.1 to 8.3 in) and the Sunda slow loris measuring 27 to 38 cm (11 to 15 in).[27] Their trunk is longer than other prosimians[38] because they have 15–16 thoracic vertebrae compared to 12–14 in other prosimians.[39] This gives them greater mobility when twisting and extending towards nearby branches.[34] Their other vertebrae include seven cervical vertebrae, six or seven lumbar vertebrae, six or seven sacral vertebrae, and seven to eleven caudal vertebrae.[39]

The eyes of slow lorises are large and have a reflective layer, called a tapetum lucidum, to help them see better at night.

Compared to galagos, which have longer legs than arms, slow lorises have arms and legs of nearly equal length.[5] Their intermembral index (ratio of arm to leg length) averages 89, indicating their forelimbs are slightly shorter than their hindlimbs.[27] Like the slender lorises, their arms are slightly longer than their body,[39] however, the extremities of slow loris are more stout.[27]

Slow lorises have a powerful grasp with both their hands and feet due to several specializations.[27][26] They can tightly grasp branches with little effort because of a special muscular arrangement in their hands and feet, where their thumb is nearly perpendicular (~180°) to the rest of the fingers and the hallux (big toe) ranges between being perpendicular to pointing slightly backwards.[26][40][5] The second digit of the hand is short compared to the other digits,[27] while on the foot, the fourth toe is the longest.[39] Their sturdy thumb helps to act like a clamp when digits three, four, and five grasp the opposite side of a tree branch.[27][5] This gives their hands and feet a pincer-like appearance.[5] Their strong grip can be held for hours without losing sensation due to the presence of a retia mirabilia (network of capillaries), a trait shared among all members of the lorisine subfamily.[28][40][5] Both slender and slow lorises have relatively short feet.[39] Like nearly all crown strepsirrhines, they have a toilet-claw or grooming claw on the second toe of each foot.[39][5]

Slow lorises have an unusually low basal metabolic rate. This may be as little as 40% of the typical value for placental mammals of their size, comparable to that of sloths. Since they consume a relatively high-calorie diet that is available year round, it has been proposed that this slow metabolism is due primarily to the need to eliminate toxic compounds from their food. For example, slow lorises can feed on Gluta bark, which can be fatal to humans.[41]

Behavior and ecology

A small 6-week-old baby clings to its mothers back as she climbs vertically through the branches
Babies cling to the mother's back.

Little is known about the social structure of slow lorises, but they generally spend most of the night foraging alone.[42][43] Slow lorises sleep during the day, usually alone but occasionally with other slow lorises.[42] There is significant overlap between the home ranges of adults, and males may have larger home ranges than females.[42][43] In the absence of direct studies of slow lorises, Simon Bearder speculated that slow loris social behavior is similar to the potto, another slow moving nocturnal primate.[44] Such a social system is distinguished by a lack of matriarchy and by factors that allow the slow loris to remain inconspicuous and minimize energy expenditure, by limiting vocal exchanges and alarm calls and by scent marking being the dominant form of communication.[44]

Slow lorises can produce a toxin which they mix with their saliva to use as protection against enemies. The toxin is similar to the allergen in cat dander.[45] Loris bites cause a painful swelling, but the toxin is mild and not fatal. Cases of human death have been due to anaphylactic shock.[46]

Slow lorises have a special network of capillaries in their hands and feet that allow them to cling to branches for hours without losing sensation.

Studies suggest that slow lorises are polygynandrous.[47] Infants are either parked on branches while their parents find food or else are carried by one of the parents.[48] Due to their long gestations, small litter sizes, low birth weights, long weaning times, and long gaps between births, slow loris populations have one of the slowest growth rates among mammals of similar size.[49]

Diet

Slow lorises are omnivores, eating insects, arthropods, small birds and reptiles, eggs, fruits, gums, nectar and other vegetation.[50][43][51][52] A 1984 study of the Sunda Slow Loris indicated that its diet consists of 71% fruit and gums and 29% insects and other animal prey.[50][43][53] A more detailed study of a different Sunda slow loris population in 2002 and 2003 showed very different dietary proportions. This study showed a diet consisting of 43.3% gum, 31.7% nectar, 22.5% fruit and just 2.5% anthropods and other animal prey.[50] The most common dietary item was nectar from flowers of the Bertram palm (Eugeissona tristis).[50] The Sunda slow loris eats insects which other predators avoid due to their repugnant taste or smell.[43] Captive slow lorises eat a wide variety of foods including bananas and other fruits, rice, dog-food, raw horse meat, insects, lizards, freshly killed chicken, mice, young hamsters and milk formula that is made by mixing instant baby food with egg and honey.[52] Captive slow loris diets may be supplemented with cod-liver oil and bone meal.[52]

Preliminary results of studies on the pygmy slow loris indicates that its diet consists primarily of gums and nectar (especially nectar from Saraca dives flowers, and that animal prey makes up 30%–40% of its diet.[50][42] However, one 2002 analysis of pygmy slow loris feces indicated that it contained 98% insect remains and just 2% plant remains.[52] The pygmy slow loris returns to the same gum feeding sites often and leaves conspicuous gouges on tree trunks when inducing the flow of exudates.[50][42] Captive pygmy slow lorises also make characteristic gouge marks in wooden substrate.[52] It is not known how the sympatric pygmy and Bengal slow loris partition their feeding niches.[50]

Slow lorises can eat while hanging upside down from a branch with both hands.[43] They spend about 20% of their nightly activities feeding.[42]

Cultural references

Beliefs about slow lorises and their use in traditional practices is deep-rooted and goes back at least 300 years, if not further based on oral traditions.[54] In the late 1800s and early 1900s, it was reported that the people from the interior of the island of Borneo believed that slow lorises were the gatekeepers for the heavens and that each person had a personal slow loris waiting for them in the afterlife. More often, however, slow lorises are used in traditional medicine or to ward off evil.[55]

In the Mondulkiri Province of Cambodia, hunters believe that lorises can heal their own broken bones immediately after falling from a branch in order to climb back up the tree, and that slow lorises have medicinal powers because they require more than one hit with a stick to die.[55] In the province of North Sumatra, the slow loris is thought to bring good luck if it is buried under a house or a road.[55][34] In the same province, slow loris body parts were used to place curses on enemies.[55] In Java, it is thought that putting a piece of its skull in a water jug would make a husband more docile and submissive, just like a slow loris in the daytime. More recently, researchers have documented the belief that the consumption of loris meat was an aphrodisiac that improves "male power." The gall bladder of the Bengal slow loris has historically been used to make ink for tattoos by the village elders in Pursat and Koh Kong Provinces of Cambodia.[55]

In Indonesia, slow lorises are called malu malu or "shy one" because they freeze and cover their face when spotted.[56]

Conservation

Main article: Conservation of slow lorises
Slow lorises are popular in the exotic pet trade, which threatens wild populations.

The two greatest threats to slow lorises are deforestation and the wildlife trade.[57] Slow lorises have lost a significant amount of habitat,[58] with habitat fragmentation isolating small populations and obstructing biological dispersal.[59] However, despite the lost habitat, their decline is most closely associated with unsustainable trade, either as exotic pets or for traditional medicine.[58] All species are listed either as "Vulnerable" or "Endangered" by the International Union for Conservation of Nature (IUCN).[59][60][61][62][63] When all five species were considered a single species, imprecise population data and their regular occurrence in Southeast Asian animal markets inaccurately suggested that slow lorises were common, resulting in a previous IUCN Red List assessment of "Least Concern" as recently as 2000.[61][64][65] Since 2007, all slow loris species are protected from commercial international trade under Appendix I.[66] Furthermore, local trade is illegal because every nation in which they occur naturally has laws protecting them.[67]

Despite their CITES Appendix I status and local legal protection, slow lorises are still threatened by both local and international trade due to problems with enforcement.[55][66] Additionally, surveys are needed to determine existing population densities and habitat viability for all species of slow loris. Connectivity between protected areas is important for slow lorises because they are not adapted to dispersing across the ground over large distances.[68]

Populations of slow loris species, such as the Bengal and Sunda slow loris, are not faring well in zoos. Of the 29 captive specimens in North American zoos in 2008, several are hybrids that cannot breed while most are past their reproductive years. The last captive birth for these species in North America was in 2001 in San Diego. Pygmy slow lorises are doing better in North American zoos. The population has grown to 74 animals between the time they were imported in the late 1980s and 2008, with most of them born at the San Diego Zoo.[57]

Wildlife trade

Prior to the 1960, the hunting of slow lorises was sustainable,[58] but due to growing demand, decreased supply, and the subsequent increased value of the marketed wildlife, slow lorises have been overexploited and are in decline.[65] With the use of modern technology, such as battery-powered search lights, slow lorises have become easier to hunt because of their eye shine.[49] Traditional medicine made from loris parts are thought to cure many diseases,[55] and the demand for this medicine from wealthy urban areas has replaced the subsistence hunting traditionally performed in poor rural areas.[65] A survey by primatologist Anna Nekaris et al. (2010) showed that these belief systems were so strong that the majority of respondents expressed reluctance to consider alternatives to loris-based medicines.[65]

Slow lorises are sold locally at street markets, but are also sold internationally over the Internet and in pet stores.[69][70] They are especially popular or trendy in Japan, particularly among women.[71][69] The reasons for their popularity, according to the Japan Wildlife Conservation Society (JWCS), are that "they're easy to keep, they don't cry, they're small, and just very cute."[71] Because of their "cuteness", videos of pet slow lorises are some of the mostly frequently watched animal videos on YouTube.[34] Despite frequent advertisements by pet shops in Japan, the World Conservation Monitoring Centre (WCMC) reported only a few dozen slow lorises were imported in 2006, suggesting frequent smuggling.[24] Slow lorises are also smuggled to China, Taiwan, Europe, the United States, and Saudi Arabia for use as pets.[71][70]

A small, young slow loris is gripped by its limbs while its front teeth are cut with fingernail cutter
Slow lorises have their front teeth cut or pulled before being sold as pets, a practice that often results in infection and death.

Even within their countries of origin, slow lorises are very popular pets,[72] particularly in Indonesia.[73] They are seen as a "living toy" for children by local people or are bought out of pity by Western tourists or expatriates. Neither local nor foreign buyers usually know anything about these primates, their endangered status, or that the trade is illegal.[74] According to 59 monthly surveys and interviews with local traders taken during late 2000s, nearly a thousand locally sourced slow lorises exchanged hands in the Medan bird market in North Sumatra.[72]

International trade usually results in a high mortality rate during transit, between 30% and 90%. Slow lorises also experience many health problems as a result of both local and international trade.[71] In order to give the impression that the primates are tame and appropriate pets for children,[75] to protect people from their potentially toxic bite,[67] or to deceive buyers into thinking the animal is a baby,[71] animal dealers either pull the front teeth with pliers or wire cutters or they cut them off with nail cutters.[74][34][72] This results in severe bleeding, which sometimes causes shock or death,[34] and frequently leads to dental infection, which is fatal in 90% of all cases.[74][75] Without their teeth, the animals are no longer able to fend for themselves in the wild, and must remain in captivity for life.[74][75] The slow lorises found in animal markets are usually underweight and malnourished, and have had their fur dyed, which complicates species identification at rescue centers.[70] As many as 95% of the slow lorises rescued from the markets die of dental infection or improper care.[75]

As part of the trade, infants are pulled prematurely from their parents, leaving them unable to remove their own urine, feces, and oily skin secretions from their fur. Slow lorises have a special network of blood vessels in their hands and feet, which makes them vulnerable to cuts when pulled from the wire cages they are kept in.[71] Slow lorises are also very stress-sensitive and do not do well in captivity. Infection, stress, pneumonia, and poor nutrition lead to high death rates among pet lorises.[74] Pet owners also fail to provide proper care because they are usually sleeping when the nocturnal pet is awake.[75]

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Literature cited

  • Bearder, Simon K. (1987). "Lorises, Bushbabies, and Tarsiers: Diverse Societies in Solitary Foragers". In Smuts, Barbara B.; Cheney, Dorothy L.; Seyfarth, Robert M.; Wrangham, Richard W.; Struhsaker, Thomas T (eds.). Primate Societies. The University of Chicago Press. ISBN 0-226-76716-7. {{cite book}}: Invalid |ref=harv (help)
  • Attention: This template ({{cite doi}}) is deprecated. To cite the publication identified by doi: 10.1007/s10764-006-9032-5, please use {{cite journal}} (if it was published in a bona fide academic journal, otherwise {{cite report}} with |doi= 10.1007/s10764-006-9032-5 instead.
  • Fitch-Snyder, H; Livingstone, K (2008). "Lorises: The Surprise Primate". ZooNooz. San Diego Zoo: 10–14. ISSN 0044-5282. {{cite journal}}: Invalid |ref=harv (help)
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  • Nekaris, N.A.I.; Bearder, S.K. (2010). "Chapter 4: The Lorisiform Primates of Asia and Mainland Africa: Diversity Shrouded in Darkness". In Campbell, C.; Fuentes, C.A.; MacKinnon, K.; Bearder, S.; Stumpf, R. (eds.). Primates in Perspective. U.S.A.: Oxford University Press. pp. 34–54. ISBN 978-0195390438. {{cite book}}: Invalid |ref=harv (help)
  • Attention: This template ({{cite doi}}) is deprecated. To cite the publication identified by doi: 10.1007/978-1-4419-1560-3_22, please use {{cite journal}} (if it was published in a bona fide academic journal, otherwise {{cite report}} with |doi= 10.1007/978-1-4419-1560-3_22 instead.
  • Nekaris, N.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.; P{anger, M.; Stumpf, R. (eds.). Primates in Perspective. U.S.A.: Oxford University Press. pp. 28–33. ISBN 978-0195171334. {{cite book}}: Invalid |ref=harv (help)
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  • Phillips, E.M.; Walker, A. (2002). "Chapter 6: Fossil lorisoids". In Hartwig, W.C. (ed.). The primate fossil record. Cambridge University Press. ISBN 978-0-521-66315-1. {{cite book}}: Invalid |ref=harv (help)
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External links

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