taxonID	type	description	language	source
CE199B17FFC1FFC4FE5A67A4F784FA66.taxon	vernacular_names	(OLD WORLD MONKEYS)	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC1FFC4FE5A67A4F784FA66.taxon	diagnosis	• Medium-sized to large monkeys with long trunk, ischial callosities, narrow nasal opening and palate, daggerlike canines, molars with two ridges, and tail lacking or short in some, long in most; males usually larger than females. • 90 - 180 cm. • Palearctic, Afrotropical, and Indo-Malayan Regions. • ‘ Tropical and subtropical forests, tropical and subtropical savanna woodland, temperate woodland, arid shrubland, and semi-desert from sea level to 4000 m.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC1FFC4FE5A67A4F784FA66.taxon	description	• 23 genera, 159 species, 269 taxa. • 18 species Critically Endangered, 40 species Endangered, 31 species Vulnerable; none Extinct since 1600	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC3FFC6FF2F675EF5BCFD31.taxon	materials_examined	Barbary Coast.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC3FFC6FF2F675EF5BCFD31.taxon	discussion	The first primate species to appear in Gesner’s 1551 Historiae Animalium and subsequently cited by Linnaeus in 1758. M. sylvanusis thought to be the earliest offshoot of the genus Macaca and is unique in being the only African macaque and the only macaque with a tail thatis vestigial or entirely lacking. Genetic studies have ascertained that the Moroccan and Algerian populations are distinct and may have split c. 1 - 6 million years ago. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC3FFC6FF2F675EF5BCFD31.taxon	distribution	Distribution. Mountainous areas in Morocco (Rif, High and Middle Atlas) and Algeria (Grande and Petite Kabylie); historically it also occurred in W Tunisia but is now extinct there. A free-ranging population occurs in Gibraltar introduced there by the British in 1740.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC3FFC6FF2F675EF5BCFD31.taxon	description	Descriptive notes. Head-body 55.7 - 63.4 cm, tail 0 - 4 - 2 - 2 mm (vestigial); weight 9 - 9 — 14 - 5 kg. Male Barbary Macaques are larger than females. Pelage coloris relatively uniform across the body but varies with age from pale buff to golden brown to dark brown in mature adults of both sexes.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC3FFC6FF2F675EF5BCFD31.taxon	biology_ecology	Habitat. A wide variety of forest habitats including cedar (Cedrus), fir (Abies), and deciduous and evergreen oak (Quercus); also thermophilous scrub and steep rocky escarpments and cliffs. The Barbary Macaque is the only non-human primate found north of the Sahara in Morocco and Algeria, with a Mediterranean climate that suffers seasonal extremes of temperature in mountainous areas.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC3FFC6FF2F675EF5BCFD31.taxon	food_feeding	Food and Feeding. Barbary Macaques are opportunistic feeders with an omnivorous diet, the content of which depends on location and season. Food items include leaves, fruit, rhizomes, seeds, lichen, animal foods, acorns, conifer seeds (cedar and fir), and fungi. They also raid agricultural crops and fruit trees in northern Morocco, and the population in the Middle Atlas has been blamed for the death of mature cedar trees because of their bark stripping behavior. Wateris a limiting factor for Barbary Macaques in some areas.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC3FFC6FF2F675EF5BCFD31.taxon	breeding	Breeding. Barbary Macaques are seasonal breeders. Females show an anogenital swelling during periovulatory periods in autumn. Females usually mate with more than one male during all stages of the ovulatory cycle. They first give birth at 4 - 6 years of age. Single infants are born in April-June every year, after a gestation of ¢. 165 days. Under captive conditions, the most fertile period of a female's life is 7 - 12 years of age. It decreases thereafter until it ceases altogether at 20 - 25 years of age. Under captive conditions, subadult males (4 - 5 - 6 - 5 years of age) have much lower reproductive success than adult males (7 - 5 - 25 years) and same-aged females. Longevity in captivity is c. 30 years for females and c ¢. 25 years for males. One of the most prominent behavioral features of the Barbary Macaque is the amount of allomaternal care provided to infants by group members. Infants in captive conditions have been recorded as spending ¢. 20 % of the day with individuals other than their mothers, particularly with adult males. Genetic testing of paternity showed that relatedness has no effect on male preference for specific infants, but infants play a role in mediating social interactions particularly among males.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC3FFC6FF2F675EF5BCFD31.taxon	activity	Activity patterns. The Barbary Macaque is diurnal, and their activity patterns are influenced by human use of their habitat and resource abundance. Depending on the time of year, groups of Barbary Macaques separate during the day to feed and come together at communal sleeping sites in trees, rocks, or caves at night. Predators include domestic dogs, Golden Jackals (Canis aureus), and Red Foxes (Vulpes vulpes).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC3FFC6FF2F675EF5BCFD31.taxon	biology_ecology	Movements, Home range and Social organization. Home ranges vary from 7 - 2 km? in the fir forest of the Moroccan Rif to 2 - 8 - 3 - 8 km? in Algeria. Home range overlap among groups can be 80 %. Overall population density estimates are 6 - 7 ind / km? in fir forest and 12 - 1 - 28 - 2 ind / km? * in the cedar forests of the Middle Atlas. Groupsize is 12 - 88 individuals. Groups of Barbary Macaques are multimale-multifemale, with rates of male emigration lower than in other macaque species. In semi-free ranging captive populations, adult Barbary Macaques are able to distinguish photographs of group members from other individuals.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC3FFC6FF2F675EF5BCFD31.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Endangered on The IUCN Red List. The Barbary Macaque has a long association with humans as evidenced by the finding of 21 mummified Barbary Macaques in an Egyptian tomb and the petrified body of a juvenile found in the Italian city of Pompeii. Serious declines in the Moroccan Middle and High Atlas populations are attributed to habitat degradation and poaching of infants for sale as pets in Morocco and Europe. The population in northern Morocco is stable or increasing and remains the best hope for the continued existence of the Barbary Macaque in the wild. Recent information on its status in Algeria is unavailable, but infant capture for the pet trade is certainly a problem there. The conservation status of the Barbary Macaqueis precarious, with less than 7000 individuals left in the wild and the main populations fragmented and distant from each other.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC3FFC7FA356602F83AF421.taxon	materials_examined	“ Ceylon. ” Corrected by J. Fooden in 1975 to India, Western Ghats, inland from Malabar coast.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC3FFC7FA356602F83AF421.taxon	discussion	M. silenus is a member of the silenus species group of macaques, with its phylogenetic sister species being M. leonina, and including M. siberu, M. nemestrina, M. pagensis, and the seven Sulawesi species. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC3FFC7FA356602F83AF421.taxon	distribution	Distribution. SW India, endemic to the hills of the Western Ghats in the states of Karnataka, Kerala, and Tamil Nadu, stretching from Anshi Ghats in the N to the Kalakkad Hills in the S, at elevations of 800 - 1300 m.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC3FFC7FA356602F83AF421.taxon	description	Descriptive notes. Head-body 51 - 61 cm (males) and 42 - 46 cm (females), tail 24 — 39 cm (males) and 25 - 32 cm (females); weight 5 - 10 kg (males) and 2 - 6 kg (females). Adult female Lion-tailed Macaques are ¢. 33 % smaller than adult males. The face is all black and surrounded by a large characteristic grayish-brown cheek and chin ruff. Fur is otherwise glossy black. The Lion-tailed Macaque is characterized by thin but strongly projecting supraorbital ridges and a flat broad nose. Eyes are closelyset, hazel, and complemented by large conspicuous eyelids contrasting with the dark face. Tailis 55 - 75 % of the head-body length and has a tuft of hair at the end, like that of a Lion (Panthera leo). Tail is normally pendulous but is carried to one side or carried up during sexual displays and carried low when frightened or to show submission.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC3FFC7FA356602F83AF421.taxon	biology_ecology	Habitat. Climax tropical evergreen and broadleaved monsoon forests (referred to regionally as “ sholas ”) with canopies reaching 30 - 50 m mostly at elevations of 100 1800 m. Lion-tailed Macaques are habitat specialists and broadly sympatric with the folivorous Nilgiri Langur (Semnopithecusjohnii). Close to the Arabian Sea, western slopes receive heavy south-west monsoons and up to 6000 mm of rain / year. Eastern slopes in the rain shadow are drier and typically have dry deciduous and scrub forests. Below 300 - 600 m, the forest grades into low-lying moist deciduous forest. Seasonally, Liontailed Macaques may enter forest at lower elevations, and they may use them more if they are not hunted. Above elevations of 1500 - 1800 m, vegetation grades into montane forest with a lower canopy. They spend most of their time in the forest canopy: 40 % at 21 - 30 m above the ground and 26 - 5 % at 11 - 20 m. In undisturbed climax forest, they are very rarely seen on the ground (just occasionally to eat mushrooms). Floristic composition differs between forests in the northern and southern parts of their distribution, probably affecting, subtly or otherwise, their diets and movements. The canopy of the northern forests (Karnataka) is abundant in Dipterocarpus (Diperocarpaceae). In the south, it is frequently dominated by Cullenia exarillata (Malvaceae), a favored food species. In some areas, Lion-tailed Macaques are able to persist in human-modified landscapes: mosaics of degraded and secondary forest patches, home gardens, and tea and cardamom plantations. Their capacity to adapt to progressively degraded forest fragments was elegantly studied by M. Singh and coworkers in the Puthuthotum coffee / tea estate, one of many on the Valparai Plateau in the Anaimalai Hills in Tamil Nadu State. Forest fragments in the region were 5 - 15,000 ha, and each was occupied by 1 - 6 groups of Lion-tailed Macaques. In 2000, a forest patch under study totaled 102 ha, of which a part (68 ha) was underplanted with cardamom and another with coffee (32 ha). The forest had been selectively logged in 1989 - 1990 and 1997 - 1998, and in many areas, non-native planted and pioneer trees predominated. Climax native species had been lost. Canopy height had decreased (19 m in 1994 to 14 m in 2000), and the canopy was severely broken (cover in 1994 was 62 % vs. 27 % in 1999). Basal area of the forest trees (area of tree trunks at breast height) dropped from 67 - 2 m? / ha to 17 - 4 m? / ha from 1994 to 1999. In 2000, c. 40 % of the area had no mature trees, and lianas and vines were almost completely absent. Two groups of Lion-tailed Macaques occupied this forest, and group size increased from 1995 to 2000 despite major changes in structure and floral composition. One group grew from 36 to 51 individuals and the other from eleven to 17 individuals. Birth rates of both groups were | infant / 2 - 4 female years (number of potentially reproductive females each year from 1996 to 2000). Lion-tailed Macaques adapted their diets and movements to these small forest patches and human modified landscapes. Compared with groups in undisturbed climax forest, they did not travel as far each day (averages of 1211 m and 734 m for the two groups compared with 1810 m for a group studied in tall forest). This was explained by a more clumped distribution of key food sources (many of them non-native and planted) of macaques in the disturbed areas. Lion-tailed Macaques wentto the ground to forage, even picking up large fallen fruits such as those of native jackfruit (Artocarpus heterophyllus, Moraceae) and Cullenia — a behavior never seen in tall forest. They also used the ground to travel through tea plantations to gain access to fruits and flowers of coffee, Lantana (Verbenaceae; invasive exotic), reach other forest patches, and even rest. Time spent in trees decreased from 95 % in 1990 to 71 % in 2000. The rate of encounters between groups increased from one encounter every eleven and a quarter hours of observation to one encounter every five and a half hours of observation in the same period. In great contrast with diets of groups in of tall-forest, nearly 40 % of the plant part of the diet involved cash crops and non-native and introduced pioneer species. The majority of the cash crop part of the diet was the mesocarp and seeds of coffee, and the incidence of coffee bushes in the forest understory resulted from macaques dispersing seeds. With the more broken canopy in the modified forest, the rate at which macaques fell out of the trees was very high (1 - 5 incidents / day). An evident effect of the forest fragmentation — the lack of any arboreal connection with other forests — was a reduced bias toward females in the adult sex ratio, because males were unable to disperse. Adult sex ratios in larger forests with multiple groups can be as high as 1: 9 - 9, but the two groups at Puthuthotum had sex ratios of 1: 4 - 6. A very large part of the remaining habitat for the Lion-tailed Macaque comprises such forest patches in human-dominated landscapes, and their capacity to adapt to them is the reason they are not on the verge of extinction.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC3FFC7FA356602F83AF421.taxon	food_feeding	Food and Feeding. The Lion-tailed Macaque is primarily frugivorous; fruit forms the major portion of its diet, supplemented with seeds, young leaves, berries, and nuts. Fruits of Artocarpus and fruits and flowers of Cullenia exarillata are important foods that are available throughout the year. Fruits of Palaquium ellipticum (Sapotaceae), Fugenia (Myrtaceae), Elaeocarpus munroii (Elacocarpaceae), Vepris biloculars (Rutaceae), and Viburnum acuminatum (Caprifoliaceae), and flowers of Loranthus (Loranthaceae) are also important in the diet in most months of the year. A number of species are important seasonally, including, for example, Syzygium mundagam (Myrtaceae), Lailsea wightiana (Lauraceae), and fruits of the palm Bentinckia condapanna (Arecaceae). Lion-tailed Macaques also eat bark, nectar, gums, and a variety of invertebrate and vertebrate prey. Fungi and lichens are eaten occasionally, and when close to villages, they eat cultivated fruits such as guava, passion fruit, coffee, and those of the exotic umbrella tree (Maesopsis eminii, Rhamnaceae), used to shade the coffee. Invertebrates and vertebrates comprise ¢. 20 % of the diet (proportion of time spent feeding on different items). The faunal component of the diet is higher in the dry season (24 %; December — - May) when fruit is scarce than in the wet season (11 %; June-November) when fruit is more abundant. They eat mostly invertebrates, including orthopterans, leaf insects (Phyllidae), and stick insects (phasmids) gleaned from the foliage and leaf litter, along with spiders picked out of their webs (communal spiders; the macaques mess with the web to scatter them and then pick them off one by one), moths and butterflies, cicadas, and dragonflies (eaten head first; wings and legs discarded), Hymenoptera and Isoptera (emerging adult forms), and caterpillars (if on a leaf, they are cajoled to a hard surface and then rubbed to remove anything extradermal — stinging hairs, stings and spines — before eating). They will eat secretions of the nest and eggs of tree frogs (Rhacophorus) and the tree frogs themselves (eaten by all ages and both sexes). Adults also eatlizards of the genera Calotes (tree lizards) and Draco (flying dragons), both in the family Agamidae. Lizards are killed by a bite to the head and eaten, head first, but without ingesting the skin and bones. Birds, nestlings, and bats are also eaten. They have been to seen to attack and severely injure the fawn of an Indian Chevrotain (Moschiola indica); it escaped and was chased, without success, by the alpha male. In certain areas, local people wrap placentas of cows in a cloth and hang them from a branch, and Lion-tailed Macaques have been seen to rip open the bundle and eat the entire placenta.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC3FFC7FA356602F83AF421.taxon	breeding	Breeding. Lion-tailed Macaques breed throughout the year but with a birth peak in February-March. The menstrual cycle is ¢. 40 days. Females are sexually mature at around four years old. Males begin to breed when they are c. 8 years old. Receptive females exhibit a conspicuous small perianal swelling. Females first give birth typically at 6 - 5 years old, and interbirth intervals have been recorded as 20 months in captivity, 23 months in a group monitored in a forest fragment in a tea plantation, and 30 months in groups in extensive rainforest habitat. The gestation period is 162 - 186 days. Neonates have brown fur and pale pink skin and are nursed for c. 12 months. Infants are weaned at 5 - 5 - 9 months. Any nipple contact post-weaning is reciprocated with punishment by the mother. Mothers invest greater parental care and interest toward male infants than female infants. Male infants are groomed twice as much as female infants. Individuals can live up to 38 years.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC3FFC7FA356602F83AF421.taxon	activity	Activity patterns. Lion-tailed Macaques are diurnal and the most arboreal of the species of macaques. Foraging for food and eating take up a little more than half of their day. They rest for ¢. 27 % of the time, travel for c. 15 %, and interact socially (mating, play, allogrooming, and agonistic interactions) for ¢. 3 - 5 %. They show no distinct seasonal patterns in activity. In cooler weather or at higher elevations, they spend more time foraging and less time resting.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC3FFC7FA356602F83AF421.taxon	biology_ecology	Movements, Home range and Social organization. Lion-tailed Macaqueslive in groups of 4 - 30 individuals, usually 10 - 20, but at least one group in a forest fragment on a tea plantation in the Anaimalai Hills had more than 50 individuals in 2001 and 84 in October 2005. This extraordinarily large group and its exceptional increase in size were attributed to high infant survival (only one of 37 infants born in 2002 - 2005 died) in the absence of hunting and with an abundance of cultivated fruits and commercial crops (e. g. coffee). Groups typically have 1 - 3 adult males and about twice the number of females, besides subadults, juveniles and infants. Ratios of adult males to adult females can be as high as 1: 9 - 9. There is a high rate of male movement among groups; females are philopatric. Male entry into other groups generally involves an aggressive takeover of the resident male rather than discretely entering the hierarchy and working one’s way up (as is typical of the Rhesus Macaque, M. mulatta, and the Japanese Macaque, M. fuscata). Male Lion-tailed Macaques in a group tend to maintain their distance from each other compared with, for example, Bonnet Macaques (M. radiata) that are friendly, huddle a lot, and spend much of their time in close proximity. Male-male relationships in Lion-tailed Macaques tend be more agonistic than affiliative, and the male dominance hierarchy is strongly linear (much more so than in Bonnet Macaques). Selective pressures hinge on the fact that male Lion-tailed Macaques breed throughout the year, which results from the relative lack of seasonality in food availability. A male, as such, is able to defend his access to females (one or two receptive at any one time), which is otherwise difficult if the breeding season is very short with all females becoming sexually receptive over a short period, as happens in Bonnet Macaques. Short studies (two months or so) have indicated home range sizes of ¢. 200 ha, but over longer periods, groups gradually change and expand their home ranges, which can extend to 500 ha or more. There is considerable overlap among home ranges of neighboring groups, but in a 500 ha home range, the central core area of ¢. 300 ha is rarely entered by other groups. Their large home ranges are related to the fact that they are selective feeders, depending on relatively few and widely dispersed food resources. Home ranges are smaller in fragmented forests next to plantations where food resources are more concentrated. Under natural circumstances, larger mammalian predators such as Leopards (Panthera pardus), Dholes (Cuon alpinus), and pythons (Python) are not a problem for Lion-Tailed Macaques because they so rarely go to the ground; however, they do mob them. Predators likely include the Indian black eagle (Ictinaetus malayensis) and the mountain hawk-eagle (Spizaetus nipalensis). Dogs are predators in humanmodified landscapes where Lion-tailed Macaques are more terrestrial.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC3FFC7FA356602F83AF421.taxon	conservation	Status and Conservation. CITES Appendix I. Classified as Endangered on The IUCN Red Last. Lion-tailed Macaques are offered the highest protection under Schedule 1, Part I, of the Indian Wildlife (Protection) Act of 1972. In the late 19 " and early 20 ™ centuries, large tracts of the forests of the Western Ghats were cleared for cultivation of rice, tea, cardamom, coffee, eucalyptus, and teak. Commercial crops such as Areca nuts and rice were even cultivated inside protected areas. Illegal settlement on reserved forests causes damage because of harvesting of non-timberforest products and the collection of firewood. Development projects such as dams (power generation and irrigation), mining and mineral exploration, road construction, and power lines threaten forests and the survival of the Lion-tailed Macaque. Today, these macaques are divided into three subpopulations. The most southerly occupies forest fragments in the Anaimalai Hills, to the south of a natural divide in the Western Ghats, the Palghat Gap in Kerala. To the north of the Palghat Gap, Lion-tailed Macaques are divided to two subpopulations due to their extirpation by hunting in the Coorg District of Karnataka. The most northerly subpopulation occurs in the region of Sirsi-Honnavara in Uttara Kannada District, Karnataka. Hunting (guns and traps) for food, traditional medicine, and the pet trade are also local but insidious threats. In 1968, it was believed that numbers of Lion-tailed Macaques in the wild did not exceed 1000 individuals. Surveys conducted in the mid- 1970 s recorded an all time low ofjust 400 macaques. In 2002, however, new and significant populations were discovered in the north ofits range, notably in the region of Sirsi-Honnavara in Karnataka, with an estimated 32 groups (790 individuals). This region is in need of protection and management. Protected areas in Karnataka are heavily hunted, and local extinction is probably inevitable if proper measures are not adopted. The total population of Lion-tailed Macaques in the wild is now estimated to be less than 3500 - 4000 individuals in 47 isolated populations, each having no more than 250 mature individuals. Mature individuals in the wild number less than 2500. About 1216 adult Lion-tailed Macaques have been reported in Kerala. In the Anamalai Hills of Tamil Nadu, ¢. 500 individuals exist in two subpopulations of 7 - 12 groups. Lion-tailed Macaques occur in a number of protected areas, including Indira Gandhi Wildlife Sanctuary and National Park, Kalakkadu Wildlife Sanctuary, Kudremukh National Park, Periyar Wildlife Sanctuary, and Silent Valley National Park.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC2FFC0FAEC6935F9D3FA53.taxon	materials_examined	“ One of the islands of the Indian Archipelago. ”	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC2FFC0FAEC6935F9D3FA53.taxon	discussion	In his 1969 review of the Sulawesi macaques, J. Fooden classified the distinct taxa as full species because he had then no evidence of intergradation. In 1980, C. P. Groves reported on a zone of intergradation between M. nigra and M. nigrescens in the downstream region of the Onggak Dumoga River, and he placed nigrescens as a subspecies of nigra. This classification was followed G. Corbet and J. Hill in their review, The Mammals of the Indomalayan Region, published in 1992. The hybrid zone: is: restricted, however, and even though M. nigra and M. nigrescens are genetically quite similar, and morphologically more similar to each other than to other species, they are distinct and since 2001, Groves has classified nigra and nigrescens as separate species. M. nigra is a member of the silenus species group, including M. siberu, M. pagensis, M. leonina, M. nemestrina, M. silenus, and the other Sulawesi species. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC2FFC0FAEC6935F9D3FA53.taxon	distribution	Distribution. N Sulawesi (E tip of the N peninsula), Manado Tua and Talise Is. Introduced into the Moluccas Archipelago (Bacan I).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC2FFC0FAEC6935F9D3FA53.taxon	description	Descriptive notes. Head-body 50 - 57 cm (males) and 44.5 - 55 cm (females), tail 1.5 - 2.5 cm; weight 10.2 - 13 kg (males) and 5.5 - 8 kg (females). Fur of the Crested Macaque is plain dark brown to blackish, as is the bare or short-haired facial skin. Infants are born with pinkish faces that turn black in the first months oflife. Hairs of the vertex are elongated (5 - 15 cm), forming an erectile crest. Ischial callosities are reniform, bright pinkish, and subdivided by a transverse furrow; in contrast, they are gray and form an elongated oval in the Gorontalo Macaque (M. nigrescens). The skull exhibits strongly developed supramaxillary ridges that are most pronounced in adult males. Posterior roots of these ridges are broadened and elevated, which gives the face of the male Crested Macaque a distinct appearance among the Sulawesi macaques (but see the Gorontalo Macaque). Male scrota and anuses have a red coloration, which has never been described for other Sulawesi macaques but may exist in at least some of them (e. g. Moor Macaque, M. maura). The Gorontalo Macaque differs from the Crested Macaque in its general body color being medium to dark brown, and its head, arms, legs, and ventral surface being black, contrasting with the color of the body. They are the same color in the Crested Macaque.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC2FFC0FAEC6935F9D3FA53.taxon	biology_ecology	Habitat. Lowland and montane primary rainforests. Due to increased pressures of habitat destruction, Crested Macaques are also found in secondary forests, actively logged forests, and grassland and cultivated land if they are surrounded by primary forest.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC2FFC0FAEC6935F9D3FA53.taxon	food_feeding	Food and Feeding. Crested Macaques have a diverse diet but mainly feed on fruits (66 % of feeding bouts) and invertebrates (32 % feeding bouts). A minor part of their diet consists of vegetative plant material and small vertebrates such as frogs, lizards, snakes, and bats. Because of the destruction of their natural habitat, Crested Macaques may invade plantations and feed on coconuts, mangos, sugar palm sap, and other crops.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC2FFC0FAEC6935F9D3FA53.taxon	breeding	Breeding. Although Crested Macaques give birth throughout the year, they show an intermediate degree of reproductive seasonality with more than three-quarters of births occurring within seven consecutive months (December — June). In the wild, females usually undergo several ovarian cycles before conceiving. Cycles are relatively long, lasting on average 39 - 6 days. In captivity, they even extend over this period if males are not available. During ovarian cycles, female Crested Macaques exhibit a pronounced swelling of their anogenital tissue, which is a good indicator of the timing of ovulation. Fully turgescent females are usually mate-guarded by high-ranking males, but they still manage to mate with other males to a minor extent. During mating, females may utter copulation calls, but the frequency and loudness of calls vary between females and between cycles of the same female. Similar to anogenital swellings, female copulation calls, if uttered, also provide a hint to the likelihood of ovulation. Male Crested Macaques may vocalize during copulation, but the function of these calls remains unknown. Similar to most of the other anthropoid primates, females do not show any post-conception mating. Infants are born after a gestation of 175 days, and females give birth only every other year. The interbirth interval is 22 months.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC2FFC0FAEC6935F9D3FA53.taxon	activity	Activity patterns. Crested Macaques are mainly terrestrial, spending 60 % of their time on the ground where they prefer to travel and rest. Most of the day, they occupy themselves moving, foraging, and feeding. Still, one-quarter of their time is spent on social activities, mainly grooming. A lot of the resting and social time takes place during midday. During extreme seasons, Crested Macaques travel less and spend more time resting and socializing on the ground, probably as an adaptation to the heat.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC2FFC0FAEC6935F9D3FA53.taxon	biology_ecology	Movements, Home range and Social organization. Crested Macaques live in multmale — multifemale, female philopatric groups, which are, in contrast to other Sulawesi macaques, quite large. Groups comprise 60 - 80 individuals (adult sex ratio: 1: 2 - 9), but they can grow up to more than 100 individuals. Smaller groups of only 25 individuals have also been observed. Home range size is 74 - 350 ha depending on group size and time of the year; home ranges are bigger during the rainy season. Home ranges of adjacent groups overlap significantly. Among macaques, Crested Macaques are among those that are the most socially tolerant, with conflicts being of lower intensity, often bidirectional and commonly reconciled. In contrast to more despotic species, female Crested Macaques often exchange affiliative interactions across matrilines. Males, however, fight fiercely over rank and can receive severe injuries. A typical demonstration of male social status is the loud call, which is emitted by all adult group males but significantly more often by high-ranking ones. Male dominance statusis also encoded in the acoustic structures of their loud calls. The red of the male’s scrotum and anus varies in intensity among individuals but also over time in the same individual. It also seems to be a sexual signal, but its precise function still remains unexplained. The extent of the red increases with male age. Males move several times between groups and also have been found roaming the forest solitarily or in small all-male groups. They emigrate from their natal group when reaching adulthood, often immediately trying to take over top dominance in neighboring groups.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC2FFC0FAEC6935F9D3FA53.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Critically Endangered on The IUCN Red List. The Crested Macaque is the most threatened of the seven species of macaques endemic to the island of Sulawesi. Crested Macaques have suffered severe hunting pressure and a widespread decline of their habitat. Consumption of monkeys and other wildlife is a longstanding tradition in ethnic groups of north-eastern Sulawesi and may have originally been sustainable. The human population in the area has substantially increased over the last decades, however, boosted by transmigration within Indonesia. This has increased hunting and led to the widespread conversion of rainforests for agricultural land and infrastructure. Legal and illegal logging and mining add to the rapid shrinking and fragmentation of natural forests, with an unknown effect on the separated small macaque populations remaining. Bushmeat hunting is not sustainable anymore and causes the rapid decline of all wildlife and, in particular, macaques. Tangkoko Batuangus Nature Reserve, although only ¢. 3600 ha, seems to be the home of the largest population of Crested Macaques within its native distribution. Nevertheless, recent surveys suggest that the population in this protected area has suffered a decline of almost 85 % since 1978. Current conservation and research activities, therefore, specifically focus on this population. A more viable population of Crested Macaques inhabits the island of Bacan, North Moluccas, where they have been introduced by humans. Given food taboos of the island’s people, the survival of this population may be better secured. The conservational value of the Bacan macaques is debatable because the island was not part of the species’ original distribution and, more importantly, the small founder population can be expected to have limited genetic variability.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC5FFC1FA2460E8FD59F912.taxon	materials_examined	Gorontalo, North Sulawesi Island, Indonesia.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC5FFC1FA2460E8FD59F912.taxon	discussion	In his 1969 review of the Sulawesi macaques, J. Fooden classified the distinct taxa as full species because he had no evidence of intergradation. In 1980, C. P. Groves reported on a zone of intergradation between M. nigra and M. nigrescens in the western tip of Bogani Nani Wartabone (Dumoga Bone) National Park in the downstream region of the Onggak Dumoga River, and he placed nigrescens as a subspecies of nigra. This classification was followed G. Corbet and J. Hill in their review The Mammals of the Indomalayan Region, published in 1992. The hybrid zone is restricted, however, and even though M. nigra and M. nigrescens are genetically quite similar and morphologically more similar to each other than to other species, they are distinct. Since 2001, Groves has classified M. nigraand M. nigrescens as separate species. Hybrids have been found with M. nigra and with M. heck: east of the Bolango River and near Bolaangitang. M. nigrescens is a member of the silenus species-group, including M. siberu, M. pagensis, M. leonina, M. nemestrina, M. silenus, and the other Sulawesi species. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC5FFC1FA2460E8FD59F912.taxon	distribution	Distribution. N Sulawesi, E of Gorontalo to Onggak Dumoga River in the N peninsula.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC5FFC1FA2460E8FD59F912.taxon	description	Descriptive notes. Head — body ¢. 60 cm (males) and ¢. 50 cm (females), tail c. 2 - 5 cm; no specific data available for body weight. Pelage of young Gorontalo Macaquesis pale brown with blackish vertex, forearms, hands, and feet. It darkens with age into dark reddish-brown to blackish, with a black median streak on the lower back. As in the Crested Macaque (M. nigra), hairs of the vertex are elongated to an erectile crest. This crest is lacking in very young individuals, but grows with age. In adults, the crestis still slightly shorter (5 - 10 cm) than in the Crested Macaque. Ischial callosities are suboval to subreniform, dark orange to dark brown and without any internal subdivision.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC5FFC1FA2460E8FD59F912.taxon	biology_ecology	Habitat. Lowland and montane rainforests up to elevations of 2000 m.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC5FFC1FA2460E8FD59F912.taxon	food_feeding	Food and Feeding. Gorontalo Macaques are more frugivorous than Crested Macaques, with fruits comprising 85 % of their diet. Arthropods and vegetative plant parts are only a minor part of their daily intake (9 % and 2: 5 %, respectively). They also raid cultivated crops (fruits, vegetables, and maize).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC5FFC1FA2460E8FD59F912.taxon	breeding	Breeding. Little is known about the reproductive biology of wild Gorontalo Macaques. A 14 month study on semi-habituated wild individuals by A. K. Kohlhaas in 1989 - 1990 revealed that females breed throughout the year, but they give birth more often during the first half of the year, similar to neighboring Crested Macaques. As for all other Sulawesi macaques, females exhibit swellings of the anogenital region, but timing of maximum turgescence with ovulation remains unclear.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC5FFC1FA2460E8FD59F912.taxon	activity	Activity patterns. Available data on sctivity patterns and strata use are derived from semi-habituated Gorontalo Macaques. They hardly come to the ground, spending more than 96 % of their time in trees. Whether or not this figure is representative for the species or an artifact of human presence needs further investigation. It has also been reported that Gorontalo Macaques spend almost half of the daytime resting, followed by engaging in social activities (mainly grooming) and moving in almost equal parts (c. 20 % each). Only 10 % oftheir daily activities consist of feeding. Increased fruit availability appears to lead to further reduction in feeding time on fruits. Similar to Crested Macaques, Gorontalo Macaques rest more during the dry season, particularly on hot days. Resting seems to center less on the midday but peaks more early and late in the day. Social activity is significantly influenced by fruit availability and temperature and, as such, decreases with increasing fruit availability and temperature.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC5FFC1FA2460E8FD59F912.taxon	biology_ecology	Movements, Home range and Social organization. Gorontalo Macaques live in multimale — multifemale, female philopatric groups, which are believed to be smaller than those of Crested Macaques. Nevertheless, because no research has been conducted on totally habituated individuals, the reliability of group counts can be questioned. The only long-term study carried out on semi-habituated animals found a mean group size of 14 individuals, but up to 65 individuals were occasionally counted in a single group, with an adult male to adult female sex ratio of 1: 1 - 7. Group sizes may have been underestimated because of group members being spread out in the dense forest. As in Crested Macaques, emigrating males often travel alone. Gorontalo Macaques are expected to be as socially tolerant as other, better studied, Sulawesi macaques, but evidence is still lacking. Agonistic behaviors appear to occur frequently, but with short duration. The severity and directionality of aggression and the frequency of reconciliation, however, remain unclear.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC5FFC1FA2460E8FD59F912.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Vulnerable on The IUCN Red List. The Gorontalo Macaqueis fully protected by Indonesian Law. Although the majority ofits distribution lies within a protected area, Bogani Nani Wartabone National Park, the population has declined in recent decades. The reason for the decline is conversion of accessible lowland rainforests, outside and in protected areas, into agricultural land and plantations. One of the affected areas is the former studysite of Kohlhaas, which has now been completely logged. There is also severe illegal hunting (market and subsistence) where macaques are not covered by a food taboo. The majority of Bogani Nani Wartabone National Park is inhabited by people of Muslim belief, and its center consists of massive, steep mountain ranges that are difficult to reach. In combination, this protects part of the population of Gorontalo Macaques relatively well. Whether Gorontalo Macaques still meet and hybridize with Heck’s Macaques (M. hecki) and Crested Macaques remains unclear but is doubtful at least for Heck’s Macaque.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC4FFC1FFFE6227F8C1F671.taxon	materials_examined	Buol, Tengah, Sulawesi, Indonesia.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC4FFC1FFFE6227F8C1F671.taxon	discussion	M. heckiis a member of the silenus species-group of macaques, including M. siberu, M. pagensis, M. leonina, M. nemestrina, M. silenus, and the other Sulawesi species. It hybridizes with M. tonkeana in an area centered on the road from Tawaeli to Toboli that crosses the base of the northern peninsula, north-east of Palu, and possibly with M. nigrescens just east Gorontalo and in the basins of the Bolango and Bone rivers. In his 1980 review, C. P. Groves considered M. hecki to be a subspecies of M. tonkeana based on inferred intergradation. This classification was followed G. Corbet and J. Hill in their review The Mammals of the Indomalayan Region, published in 1992. The hecki-tonkeana hybrid zone was subsequently studied by E. Bynum and colleagues and found to be restricted (c. 15 km by 7 - 5 km) and old, dating back to at least to 1900. This indicates that it is a zone of secondary contact and that there are barriers (prezygotic or postzygotic) to full introgression. In 2001, Groves listed heck: as a distinct species. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC4FFC1FFFE6227F8C1F671.taxon	distribution	Distribution. NW Sulawesi from the base of the N peninsula (Isthmus of Palu) NE to just E of Gorontalo.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC4FFC1FFFE6227F8C1F671.taxon	description	Descriptive notes. Head — body 58 - 68 cm (males) and 50 - 57 cm (females), tail 2 - 3 cm (males) and 2 - 3 cm (females); weight 8 - 10 kg (males) and 7 - 8 kg (females). Heck’s Macaque is black, with a brownish underside. Forearms are dark brown, and shanks are gray to light brown (i. e. paler than the trunk). Crown hair is lengthened but does not form a distinct crest, and cheeks are lighter colored and without elongated whiskers. Tail is very short and stubby, although not rudimentary. Ischial callosities are gray to yellow and kidney-shaped, with only a partial transverse furrow across them. Comparing the Gorontalo Macaque (M. nigrescens) with Heck’s Macaque, the Gorontalo Macaque has a baboon-like face, an elongated narrow crest, blackish limbs contrasting with a brown body, and gray ischial callosities. Heck’s Macaque has a short broad face and a short broad, somewhat upstanding crest, on the crown. It is black with strongly contrasting brown shanks (extending up the back of the thighs), weakly contrasting dark brown arms and underparts, and yellow, reniform, and only partially separated callosities. Comparing the Tonkean Macaque (M. tonkeana) with Heck’s Macaque, forearms and shanks of hindlimbs of the Tonkean Macaque are black like the body, underside is no paler than the upper side (it is in Heck’s Macaque), cheek whiskers have a pale grayish or brownish tint (not in Heck’s Macaque), rump patch is more clearly marked and bushy (not so in Heck’s Macaque), and ischial callosities are oval and orange-colored (similar to the Moor Macaque, M. maura, and the Booted Macaque, M. ochreata).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC4FFC1FFFE6227F8C1F671.taxon	biology_ecology	Habitat. Undisturbed or only lightly disturbed primary rainforest at elevations from close to sea level to 1600 - 1800 m. Seasons are not pronounced in these areas, with only one or two drier months each year and annual rainfall of 2500 - 3500 mm (higher in the mountainous regions). The extent to which Heck’s Macaque can thrive in degraded and secondary forest is not known.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC4FFC1FFFE6227F8C1F671.taxon	food_feeding	Food and Feeding. The three macaque species of the northern peninsula, including Heck’s Macaque, are all principally frugivores / insectivores and are less folivorous than species to the south. When near human settlements, Sulawesi macaques raid subsistence and cash crops such as maize, yams, cassava, papaya, banana, and cacao, and are persecuted for doing so.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC4FFC1FFFE6227F8C1F671.taxon	breeding	Breeding. Reproductive cycle of female Heck’s Macaquesis ¢. 36 days. As in all Sulawesi macaques, females exhibit a pronounced, bright pink to red sexual swelling during the periovulatory period. A single offspring is born after a gestation of 174 - 196 days. Young are nursed for c ¢. 12 months and are sexually mature at c. 4 years of age. Sexual patterns are broadly similar among Sulawesi macaques. The only behavioral patterns that appear to be specific to each taxon are their loud calls or chirp barks.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC4FFC1FFFE6227F8C1F671.taxon	activity	Activity patterns. Heck’s Macaques are diurnal and largely arboreal. The extent to which they become terrestrial, a trend for all macaques that are losing their forests, is unknown.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC4FFC1FFFE6227F8C1F671.taxon	biology_ecology	Movements, Home range and Social organization. Heck’s Macaque is one of the least studied of the Sulawesi macaques. It lives in multimale — multifemale groups averaging 20 individuals (range 7 - 30) and probably in large home ranges of more than a 100 ha. As in all Sulawesi macaques (and typical for the genus), males disperse and females are philopatric. Densities are 20 - 50 ind / km?.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC4FFC1FFFE6227F8C1F671.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Vulnerable on The IUCN Red List. Heck’s Macaque is fully protected by Indonesian Law. It is threatened mainly by loss of habitat, which is increasingly fragmented. The destruction of Sulawesi forests accelerated in the last 40 years, as it has for tropical forests worldwide; commercial logging, Indonesia’s transmigration program, the government-subsidized cash crop industry, and human population growth are all to blame. A report by C. Cannon and colleagues in 2007 reported that 80 % of Sulawesi forests had been destroyed. Heck’s Macaque is hunted in some areas and persecuted as a crop pest. The fearis that intensive hunting for commercial trade within the ranges of the Crested Macaque (M. nigra) and the Gorontalo Macaque, which has resulted in their severely reduced numbers and endangered status, will move south in the northern peninsula into the range of Heck’s Macaque. In 1994, it was estimated that the total population was c. 100,000, but it has certainly declined since then. Forest remains in inaccessible areas, but lowlands in the southern part of its distribution and coastal areas of the northern peninsula, in particular, have been largely converted to agriculture. The suitability of forests for Heck’s Macaques at higher elevations is unknown. In 1999, the lack of strictly protected areas in the range of Heck’s Macaque was seen as the biggest threat to its survival. Protection then was only afforded by a single nature reserve, Panua, of insufficient size (1500 ha), badly degraded and harboring low numbers of macaques. The Paguyaman River catchment was identified as of conservation value for Heck’s Macaque. Since 1999, Nantu Wildlife Reserve and a number of other reserves have been created that protect, or should protect, Heck’s Macaque: Dolangan, Mas Popaya Raja, Pangi Binanga, Pinjam / Tanjung Mantop, and Tangale reserves.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC4FFC2FAC66CC6F752FC22.taxon	materials_examined	Indonesia, Sulawesi Tengah.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC4FFC2FAC66CC6F752FC22.taxon	discussion	M. tonkeana hybridizes with M. ochreata, M. maura, and M. hecki in areas where their distributions overlap. Genetic studies have found little evidence of substantial gene flow between parental species outside of the narrow hybrid zones. Based on morphometric and dermatoglyphic (fingerprint) data, eastern populations (east of Bongka River, including those found on the Togian Islands) were proposed to constitute a separate species, M. togeanus, the “ Balantak Macaque. ” Subsequent molecular research, however, did not identify any fixed molecular markers in nuclear DNA that would support a separate species status for these populations. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC4FFC2FAC66CC6F752FC22.taxon	distribution	Distribution. C Sulawesi (S to Latimojong, SW to the base of the Toraja highlands at the Tempe Depression, SE toward, but not at, the lakes region of the SE peninsula, and NW to the isthmus between Palu and Parigi) and Togian Is.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC4FFC2FAC66CC6F752FC22.taxon	description	Descriptive notes. Head — body 58 - 68 cm (males) and 50 - 57 cm (females), tail 4 - 7 cm (males) and 3 - 6 cm (females); weight 10 - 12 kg (males) and 8: 6 - 10.4 kg (females). The Tonkean Macaque has black pelage, with white hair on the rump, dorsal parts of hindlimbs, and parts of the ventrum. Cheek tufts are prominent and range in color from pale brownish-gray to a pale brownish-yellow. Supraorbital ridges are strongly developed. There is a sagittal crest, and the muzzle is elongated, with lateral ridges. Ischial callosities are oval-shaped and orange-toned, and they have recently been characterized (along with those of the Moor Macaque, M. maura) as Type 1: oval without bending. Some show graying or whitening on their face, particularly around their eyes, head, and back, or entire bodies.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC4FFC2FAC66CC6F752FC22.taxon	biology_ecology	Habitat. Primary and secondary tropical rainforest at predominantly hill (400 - 850 m) and upland (850 - 1500 m) elevations, but also in montane forest (1500 - 2500 m). Tonkean Macaques can also exist in anthropogenically altered environments, incorporating agricultural areas into their usable habitat. Rainfall and the availability of forest fruit show fairly uniform temporal distributions in Lore Lindu National Park. Figs (Ficus, Moraceae), an important food resource for much of the Sulawesi fauna, are very abundant in Lore Lindu (34 - 8 figs / ha) compared with other Indonesian forests (6 - 6 figs / ha, East Kalimantan Province and 7 - 10 figs / ha, North Sulawesi Province). Tonkean Macaques are frequently found in association with the endemic yellow-billed malkoha (Phaenicophaeus calyorhynchus). Malkohas benefit from insect prey flushed from the forest canopy by moving macaques.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC4FFC2FAC66CC6F752FC22.taxon	food_feeding	Food and Feeding. Tonkean Macaques have a diverse diet (more than 55 plant species). Ripe fruit makes up the largest part (c. 70 - 80 %). Key fruit species include Ficus (Moraceae), Elmernillia tsiampacca (Magnoliaceae), Artocarpus (Moraceae), Pandanus (Pandanaceae), Pangium edule (Salicaceae), and, particularly in anthropogenically altered areas, the sugar palm (Arenga pinnata, Arecaceae). Insects, young leaves, shoots, and stalks (13 - 18 % ofdiet) are also frequently consumed. Groupsliving in close proximity to agricultural land incorporate cultivated crops into their diet (e. g. bananas, papaya, maize, cacao, sweet potato, and legumes).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC4FFC2FAC66CC6F752FC22.taxon	breeding	Breeding. Reproductive data come from captive populations of Tonkean Macaques. Females begin cycling at 4 - 5 years old, have a mean cycle length of 37 - 41 days, and exhibit a pronounced, bright pink to red sexual swelling during the periovulatory period. In young females, the only area that reddens and swells is between the anus and the base of the tail. As females age, the swelling becomes larger and more pronounced. A distinctive feature of Tonkean Macaques is the appearance of two tumescent lumps on the rump that develop under the fur on both sides of the spine. Mean duration of genital swelling is 13 days. Consortships begin at the time of swelling and last 5 - 10 days. A single offspring is born after a mean gestation of 173 days. In the wild, some female Tonkean Macaques have a one-year interbirth interval.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC4FFC2FAC66CC6F752FC22.taxon	activity	Activity patterns. Tonkean Macaques are diurnal, arboreal, and terrestrial. Increased terrestriality (more than 20 % of behavioral records) and more time devoted to foraging were observed in disturbed habitat. Groups spend, on average, 34 % of their day resting, 33 % moving, 23 % foraging and feeding, and 10 % socializing. Several cases of tool use have been observed in captive groups of Tonkean Macaques (e. g. using sticks to obtain food and leaning a log against a wall to climb on it).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC4FFC2FAC66CC6F752FC22.taxon	biology_ecology	Movements, Home range and Social organization. Tonkean Macaques live in multimale-multifemale groups. Mean group size is 24 individuals (range 6 - 35), with an adult male to adult female sex ratio of 1: 0 - 6 - 1 - 3. Subgrouping has been observed. Females form linear dominance hierarchies, but they show relaxed relationships characterized by low levels of intense aggression, bidirectionality when agonistic behavior occurs (either female in a dyad may initiate the interaction), frequent post-conflict affiliative contacts, and an overall high degree of social tolerance (e. g. feeding together). Adult males also show social tolerance as exemplified by low levels of agonistic behavior and frequent grooming interactions. Adult males (typically the highest ranked) emit a birdlike vocalization called a “ loud call. ” These calls appear to function primarily to influence intragroup cohesion, i. e. to communicate information with regard to location and movement within the social group. Home range size is 45 - 143 ha, and the ranges of neighboring groups often overlap. Tonkean Macaques are flexible in response to habitat alteration by intensively using particular areas within their home range where known resources are present and predictably available (e. g. A. pinnata palms). Individuals also increase their home range size under such conditions. Mean daily movement is 1175 m. Densities of Tonkean Macaques are 20 - 67 ind / km? across their distribution.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC4FFC2FAC66CC6F752FC22.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Vulnerable on The IUCN Red List. Overall population size of the Tonkean Macaque is ¢. 150,000 individuals. It has the largest distribution of all Sulawesi macaques. It is known to occurin at least three protected areas: Lore Lindu National Park, Morowali Nature Reserve, and Faruhumpenai Nature Reserve, totaling at least 544,000 ha of protected area. Separate conservation management of eastern and western populations of Tonkean Macaques has been suggested, given their genetic distinctiveness. The greatest threat is large-scale conversion of forest for commercial logging and the development of cash-crop plantations, such as oil palm and cacao. Recent reports indicate that 80 % of Sulawesi’s forests have been altered and destroyed, and only 30 % of the remaining forest is in good condition (i. e. forest canopy unbroken by large clearings and with only scattered signs of human activity). A key concern is that much of the remaining forest is montane (e. g. 90 % in Lore Lindu National Park is montane forest ranging up to elevations of 2355 m), and itis not yet certain if forest above 2000 m provides habitat that can sustain Tonkean Macaques throughout the year. Tonkean Macaques are hunted for food in some areas, particularly in non-Muslim areas where consumption of macaques is not taboo. There is also evidence of hunting in Central Sulawesi for local consumption and bushmeat markets in Manado, North Sulawesi. In many areas, and regardless ofreligion, Tonkean Macaques are poisoned or trapped as crop pests. They are also often trapped to sell as pets. Pet ownership is linked to disease transmission between macaques and people. For example, pet Tonkean Macaques from the eastern peninsula of Central Sulawesi showed evidence of exposure (seropositivity for antibodies) to endemic human pathogens, including measles, influenza A, and parainfluenza 1, 2, and 3. The fact that the same human pathogens have been detected in wild populations suggests that pet macaques may act as vectors from human to wild populations because wild macaques are often attracted to pet animals (e. g. when females are receptive), therefore putting wild populations at risk. Human-macaque folklore in the Lindu highlands serves to protect local macaques from retaliation by farmers bacause of crop raiding. It remains unknown, however, at what threshold crop losses will no longer be tolerated, and social taboos, and the conservation outcomes they afford, will be abandoned.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC7FFC3FA2E6132FCE5F8E4.taxon	materials_examined	Sulawesi Selatan, Indonesia.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC7FFC3FA2E6132FCE5F8E4.taxon	discussion	Along with other Sulawesi macaques, M. maura is a member of the silenus species group, including M. siberu, M. pagensis, M. leonina, M. nemestrina, and M. silenus. M. maura hybridizes with M. tonkeana in areas where their distributions overlap (base of the Toraja Highlands; Tempe Depression). Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC7FFC3FA2E6132FCE5F8E4.taxon	distribution	Distribution. SW Sulawesi (from Bontobahari at the tip of the SW peninsula N to Tempe Depression).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC7FFC3FA2E6132FCE5F8E4.taxon	description	Descriptive notes. Head — body 55 - 69 cm (males) and 46 - 59 cm (females), tail 5 - 7 cm (males) and 2 - 4 cm (females); weight 8.2 - 10 kg (males) and 3.8 - 7.6 kg (females). The Moor Macaque is primarily a brown or brown-black species noted for its short, dark, angular face, extremely prominent brow ridges, and flat crown hair. Rump patch is small and brownish-gray. Ischial callosities are oval shaped and orange toned, and they have been recently been characterized (along with the Tonkean Macaque, M. tonkeana) as Type 1: oval without bending. Some Moor Macaques show whitening on their face, particularly around their eyes, head, back, or their entire bodies.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC7FFC3FA2E6132FCE5F8E4.taxon	biology_ecology	Habitat. Moist deciduous forest (up to 2000 m above sea level), mosaics of forest with grasslands resulting from habitat conversion, and areas of cultivation surrounded by primary and secondary forests. A large part of the remaining population of Moor Macaques is likely confined to karst (limestone) forest. The tower karsts in southwestern Sulawesi are 150 - 300 m tall, with many fruit trees growing on top that are favored by Moor Macaques. Karst forests in Bantimurung-Bulusaraung National Park have considerable seasonal variation in rainfall, with a pronounced dry season in May-September. Moor Macaques are frequently found in association with endemic yellowbilled malkohas (Phaenicophaeus calyorhynchus). Malkohas benefit from insect prey that is flushed out of the canopy by the macaques’ movements. Figs (Ficus, Moraceae), an important food resource for many Sulawesi mammals and birds, are very abundant in Karaenta Nature Reserve (23 - 3 fig trees / ha) compared with other Indonesian forests (6 - 6 fig trees / ha, East Kalimantan Province and 7 - 10 fig trees / ha, North Sulawesi Province).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC7FFC3FA2E6132FCE5F8E4.taxon	food_feeding	Food and Feeding. Moor Macaques are primarily frugivorous, feeding largely on fruits of Ficus, rao (Dracontomelon mangiferum, Anacardiaceae), pangi (Pangium edule, Salicaceae), bakan (Litsea firma, Lauraceae), and the “ Binkuru ” tree. Insects and leaves from herbaceous vegetation are also frequently eaten (the latter particularly during the dry season). Rice, cacao, and maize are also eaten where home ranges of Moor Macaques neighbor agricultural land.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC7FFC3FA2E6132FCE5F8E4.taxon	breeding	Breeding. Reproductive cycles of female Moor Macaques are c. 36 days. Females exhibit regular repeated sexual swellings. These swellings are large and pink, extend above the ischial callosities, and last for an average of twelve days. Females first begin to show sexual swellings at 4 - 6 years old. They show moderate birth seasonality, with greater than 30 % of births occurring within a 3 month period. Females first give birth at 6 - 7 years old. A single offspring is born after a gestation of 174 - 196 days. Interbirth intervals are c. 1 - 5 - 2 years. Newborns have a dusky-gray face with pale upper eyelids and black fur. Individuals may live up to 28 years.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC7FFC3FA2E6132FCE5F8E4.taxon	activity	Activity patterns. Moor Macaques are diurnal and both arboreal and terrestrial. A group in Bantimurung-Bulusaraung National Park traveled on the ground and climbed trees and karst formation when foraging. This group spent, on average, 40 - 8 % of their time resting, 30 % moving, and 29 - 2 % feeding.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC7FFC3FA2E6132FCE5F8E4.taxon	biology_ecology	Movements, Home range and Social organization. The Moor Macaque lives in multimale-multifemale groups. Group sizes are 15 - 40 individuals, with an adult sex ratio of males to females of 1: 0 - 5 - 2 - 7. Males emigrate from their natal groups at 7 - 9 years of age. Females form linear dominance hierarchies, but they have relaxed relationships characterized by low levels of intense aggression, bidirectionality when agonistic behavior occurs (either female in a dyad may initiate the interaction), and frequent post-conflict affiliative contacts. Social tolerance among adult males has also been reported. Adult males (typically the highest ranked) emit a birdlike vocalization called a “ loud call. ” These calls are given when group members are dispersed, following stimuli such as agonistic interactions or vocalizations from other group members, but not during intergroup interactions. Home range size is 20 - 30 ha, and ranges often overlap with those of neighboring groups. Data from the group in Bantimurung-Bulusaraung National Park indicate that intergroup encounters occur, on average, every 28 - 8 hours. Males, rather than females, appear to be the primary participants. Presence of sexually receptive females is not related to frequency of intergroup encounters. Densities are 25 - 50 ind / km? ®. The well-studied population that lives in Karaenta Nature Reserve in Bantimurung-Bulusaraung National Park reached a density of 70 ind / km? * by 1998.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC7FFC3FA2E6132FCE5F8E4.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Endangered on The IUCN Red List. The major threat to the Moor Macaque is habitat disturbance, fragmentation, and overall loss of habitat. About 80 % of Sulawesi’s forests have been altered or destroyed, and of the remaining, only 30 % is in good condition (i. e. continuous forest with minimal signs of human activity). These factors are largely due to a growing human population on the island, with south-western Sulawesi being the region with the second highest human density of 129 people / km *. Because the Moor Macaque raids crops, it is also threatened by farmer retaliation. Farmers poison, trap or shoot crop raiders, and often sell live captures as pets. In 1992, the total population of Moor Macaques was ¢. 3000 - 5000 individuals, down from ¢. 56,000 individuals in 1983. Survey methods used for these population estimates differed, however, and the level of decline over three generations was more likely 50 - 80 %, or even greater than 80 %. The current population size is unknown. Although it is likely that an observer can encounter Moor Macaques in various parts of the original distribution, populations are extremely fragmented and probably largely confined to karst landscapes within protected areas. The primary protected area where the Moor Macaque occurs is Bantimurung-Bulsaraung National Park, but it is believed to occur in only 8700 ha of the 43,500 ha Park. In the 1990 s, Gunung Lompobatang was identified as a potential forest refuge and was recommended for upgrade to IUCN Category I or II (i. e. nature reserve or national park status). To date, this forest remains unprotected. There is an urgent need to address cement mining of karst areas because, heretofore, the Moor Macaque and other species have been protected by the inaccessibility of these formations.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC6FFC3FFEA6271F9EEF7AD.taxon	materials_examined	type locality unknown.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC6FFC3FFEA6271F9EEF7AD.taxon	discussion	M. ochreata is a member of the silenus species group of macaques, including M. siberu, M. pagensis, M. leonina, M. nemestrina, M. silenus, and the other Sulawesi species. The form brunnescens was listed as a species byJ. Fooden in his 1969 review, but he also found it to be the least differentiated, being very similar to its neighbor, ochreata, on mainland south-eastern Sulawesi. In their review, The Mammals of the Indomalayan Region, published in 1992, G. Corbet and J. Hill considered brunnescens to be a junior synonym of M. ochreata. It has been separated from ochreata by a maximum of only ¢. 10,000 years. Two subspecies recognized.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC6FFC3FFEA6271F9EEF7AD.taxon	distribution	Subspecies and Distribution. M. o. ochreataOgilby, 1841 — SESulawesi, throughtheentireSEpeninsula, extendingtotheNofthelakesregion; intheE, theborderbetweenthissubspeciesandtheTonkeanMacaque (M. tonkeana) istheLaRiver, whileintheWitextendsalongthecoast, acrosstheKaraenaRiverinitslowercourse, butnotreachingintouplandsfurtherinland. M. o. brunnescens Matschie, 1901 — Muna and Buton Is, and possibly the neighboring islet of Pulau Labuan Blanda, off the coast of Sulawesi; it may no longer occur on Muna because it is almost completely deforested.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC6FFC3FFEA6271F9EEF7AD.taxon	description	Descriptive notes. Head-body 48 - 59 cm (males) and 40 - 48 cm (females), tail 4 - 6 cm (males) and 3 - 5 cm (females); weight 6 - 9 kg (males) and 3 - 5 kg (females). The Booted Macaqueis blackish or brown above, with paler forearms, shanks, and rump. Hairs on the crown are darker than surrounding fur and stand erect, forming a circular cap. Tail is button-like, and ischial callosities are oval and bend outward (Type 2). The nominate form ochreata is mostly blackish, with forearms, shanks, and rump ocherousgray to whitish. The “ Buton Macaque ” (M. o. brunnescens) is not as dark as the mainland nominate subspecies, being generally brown, with forearms, shanks, and rump a lighter shade, and has a shorter face.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC6FFC3FFEA6271F9EEF7AD.taxon	biology_ecology	Habitat. Nominate form found in mangrove and coastal forests, savanna habitat (Rawa Aopa Watumohai National Park), and steep upland tropical forest at elevations up to 800 m (Faruhumpenai Nature Reserve). The Buton Macaque occupies lowland and hill forest on the island of Buton.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC6FFC3FFEA6271F9EEF7AD.taxon	food_feeding	Food and Feeding. Systematic feeding data come from short-term studies of the Buton Macaque: 66 % fruit (including crops), 20 % stems and flowers, 12 % leaves, and 2 % invertebrates. Wild fruit consumed include Ficus (Moraceae) and kapok (Ceiba pentandra, Malvaceae). Cultivated crops (e. g. cacao, sweet potato, bananas, and maize) comprised 24 % of food items eaten. The most common cultivated crop raided is sweet potato. Near Faruhumpenai Nature Reserve, the nominate subspecies is known to raid cacao plantations.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC6FFC3FFEA6271F9EEF7AD.taxon	breeding	Breeding. Female Booted Macaques exhibit a sexual swelling during their periovulatory period. They are assumed to have a c. 36 day reproductive cycle and a gestation of c ¢. 24 weeks, as has been found for other Sulawesi macaques. Females and males are sexually mature at about five and seven years old, respectively.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC6FFC3FFEA6271F9EEF7AD.taxon	activity	Activity patterns. Booted Macaques are diurnal, arboreal, and terrestrial. Buton Macaques spend about the same amount of time in trees as they do on the ground. The level of terrestriality depends on habitat type; they are more terrestrial in areas disturbed by people. Activity budget data come from the Buton Macaque. It spends the majority of its time traveling between feeding sites (43 %) and foraging and feeding (21 %), followed by resting (17 %) and grooming (10 5 %). Crop-raiding groups spend more time grooming and resting than forest-based groups, which spend more time traveling. Those that do raid crops spend ¢. 32 - 38 % of their day engaged in this activity.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC6FFC3FFEA6271F9EEF7AD.taxon	biology_ecology	Movements, Home range and Social organization. Booted Macaques live in multimale-multifemale groups. Group size is 12 - 28 individuals for the nominate subspecies and 20 - 40 individuals (mean 32) for the Buton Macaque. The Buton Macaque has an adult male to female sex ratio of 1: 2. Larger groups of Buton Macaques have been observed breaking up into smaller subgroups. The mean home range size of Buton Macaques is 62 ha. Population densities of the nominate subspecies are 0 - 97 - 2 groups / km?. The mean density of three study groups on Buton was 55 - 3 ind / km? *.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC6FFC3FFEA6271F9EEF7AD.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Vulnerable on The IUCN Red List, including both subspecies. Both subspecies of the Booted Macaque are primarily threatened by conversion of habitat to human uses. While they are not typically killed for food, poisoned bait is used against crop-raiding macaques by local cacao farmers. The nominate subspecies is considered Vulnerable because of the projected increase in oil palm, cacao, and cotton plantations and human settlement. It is known to occur in at least five protected areas: Tanjung Batikolo Wildlife Reserve, Tanjung Peropa Game Reserve, Rawa Aopa Watomahai National Park, and Padang Mata Osu and Faruhumpenai nature reserves. The population size of the Booted Macaque is ¢. 71,000 individuals. On Buton Island, the Buton Macaque is considered Vulnerable because of the projected increase in teak plantations, illegal logging, and human settlements. In addition, the extent of its occurrence is less than 20,000 km?, and this area is severely fragmented and continuing to diminish. The rate of deforestation in the area where the Buton Macaque is well studied is ¢. 10 % / year and increasing annually. The Buton Macaque is known to occur in three protected areas: Lambusango, Buton Utara, Napabalano reserves. Its population size ¢. 40,000 individuals.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC6FFCCFAF26D89FD0DF8EB.taxon	materials_examined	Indonesia, South Pagai Island.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC6FFCCFAF26D89FD0DF8EB.taxon	discussion	M. pagensis was considered a subspecies of M. nemestrina until additional study led to the conclusion that it was a separate species. It was later considered a separate species from M. siberu based on deep genetic and morphological divergence. M. pagensisis a member of the silenus species group of macaques, including M. siberu, M. leonina, M. nemestrina, M. silenus, and the Sulawesi species. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC6FFCCFAF26D89FD0DF8EB.taxon	distribution	Distribution. Mentawai Archipelago (Sipora, North Pagai, and South Pagai, and on the Sinakak islet off the E coast of South Pagai).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC6FFCCFAF26D89FD0DF8EB.taxon	description	Descriptive notes. Head — body 53 cm (males) and 43 - 46 cm (females), tail 13 - 16 cm (males) and 10 - 13 cm (females); weight 6 - 9 kg (males) and 4.5 - 6 kg (females). The Pagai Macaque is similar to the pig-tailed macaques, except for its almost hairless tail and darker dorsal coloration. Fur is a non-agouti chocolate brown above, with a paler chocolate brown on the ventral side. Neck patches are ocherous buff, and cheek patches are absent. Tail is less than 35 % of head-body length and is hairless except at the base. The Pagai Macaque is similar to the Siberut Macaque (M. siberu), but its coloration is lighter and its facial structure is not as broad.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC6FFCCFAF26D89FD0DF8EB.taxon	biology_ecology	Habitat. Primary and secondary forest areas, as well as coconut groves near villages. Densities of Pagai Macaques do not differ significantly across most forest types. Nevertheless, they live at higher densities in logged forest than in unlogged forest, and their highest density is in forest logged 20 years ago. The Pagai Macaque was the most abundant primate species detected along line transect throughout the Pagai Islands. Mentawai forests are ever-wet rainforests, with annual rainfall of up to 4000 mm.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC6FFCCFAF26D89FD0DF8EB.taxon	food_feeding	Food and Feeding. The Pagai Macaque is primarily frugivorous, presumably similar to the Siberut Macaque. It eats coconuts, but only adult males are able to open them.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC6FFCCFAF26D89FD0DF8EB.taxon	breeding	Breeding. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC6FFCCFAF26D89FD0DF8EB.taxon	activity	Activity patterns. The Pagai Macaque is diurnal and terrestrial and arboreal. Vocalizations are heard in the early morning hours.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC6FFCCFAF26D89FD0DF8EB.taxon	biology_ecology	Movements, Home range and Social organization. Pagai Macaques spend most of their time in the lower levels of the forest and on the ground. Group size is 5 - 25 individuals, and the largest group observed had two adult males. Groups typically split into smaller foraging groups during the day, but they move as one large group through coconut groves and primary forest. Pagai Macaques interact with two endemic colobines, the Mentawai Langur (Presbytis potenziani) and the Pig-tailed Langur (Simias concolor). Home range size is not known.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC6FFCCFAF26D89FD0DF8EB.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Critically Endangered on The IUCN Red List. In recent years, logging has decimated the habitat of the Pagai Macaque, and hunting has increased along with a burgeoning human population and because of improved access to remote areas due to logging roads and tracks. Bows and arrows have been replaced with rifles, and local rituals and taboos that formerly regulated hunting have been lost. Pagai Macaques are often poisoned or trapped and killed as crop pests. The most recent estimates of population density suggest 7 - 12 ind / km? in suitable habitat on the Pagai Islands, giving a total of ¢. 2100 - 3700 individuals (down from 15,000 in 1980). The pet trade may also be a threat to Pagai Macaques; they are kept and sold as pets on the Mentawai Islands. Mentawai macaque meat is considered unpalatable, but they are one of the easiest primates to catch because they can be caught by setting traps. They are frequently trapped because they are crop pests. While habitat disturbance appears to positively affect densities of Pagai Macaques, they are found in lower densities near human settlements. The Pagai Macaque is not found in any protected areas, but it does occur in some areas that have been suggested for protection (including Betumonga Research Area and the Sinakak islet, both of which were reportedly logged in recent years). All populations of Pagai Macaques are in urgent need of protective measures. It has been kept in zoos, but there is no viable captive population. Only 10 - 15 % of the original forest cover remains on Sipora. An action plan, published in 2006, suggested the following conservation actions: protection of areas in the Pagai Islands by cooperating with a logging corporation that has practiced sustainable logging there since 1971; conservation education, especially regarding hunting; and the development of alternative economic models for the local people to reduce the likelihood of selling their lands to logging companies.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC9FFCCFFD5624FF654F9FF.taxon	materials_examined	Indonesia, Siberut Island.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC9FFCCFFD5624FF654F9FF.taxon	discussion	M. siberu was first described by Fuentes and Olson in 1995 as a subspecies of M. pagensis. In 2002, A. Kitchener and C. P. Groves reviewed the taxonomy of the species and concluded that it was sufficiently distinct to warrant species status as M. siberu. M. siberu is a member of the silenus species group of macaques, including M. pagensis, M. leonina, M. nemestrina, M. silenus, and the Sulawesi species. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC9FFCCFFD5624FF654F9FF.taxon	distribution	Distribution. Mentawai Archipelago (Siberut I).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC9FFCCFFD5624FF654F9FF.taxon	description	Descriptive notes. Head-body 47 - 48 cm (males) and 40 - 45 cm (females), tail 13 — 16 cm (males) and 10 - 13 cm (females); weight 6 - 9 kg (males) and 4.5 - 6 kg (females). The Siberut Macaque is similar to the Pagai Macaque (M. pagensis), but it has ajet-black back and a white to silvery-white underside. Cheek patches of the Siberut Macaque are white, and it lacks neck patches as are found on the Pagai Macaque. Ischial callosities are light-colored. Tail is hairless except at the base and much shorter than that of the Pagai Macaque (less than 20 % of head-body length). Facial structure of the Siberut Macaque is broader than in the Pagai Macaque.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC9FFCCFFD5624FF654F9FF.taxon	biology_ecology	Habitat. Nearly all types of forest, including primary dipterocarp forest, primary mixed forest, mangrove, secondary forest, riparian forest, logged forest, and nipa palm groves. Siberut Macaques strongly prefer primary forest. Mentawai forests are ever-wet rainforests, with annual rainfall of up to 4000 mm.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC9FFCCFFD5624FF654F9FF.taxon	food_feeding	Food and Feeding. Ripe fruits are predominant in the diet of the Siberut Macaque (c. 75 %); they also eat arthropods, leaves, fungus, and bark.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC9FFCCFFD5624FF654F9FF.taxon	breeding	Breeding. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC9FFCCFFD5624FF654F9FF.taxon	activity	Activity patterns. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC9FFCCFFD5624FF654F9FF.taxon	biology_ecology	Movements, Home range and Social organization. Group cohesion of Siberut Macaques appears to be low; they often split into smaller foraging groups during the day. Travel (2 - 3 km / day) is mostly terrestrial, they feed in the canopy, and rest in the middle forest layers. Siberut Macaques spend most of their time in the middle layer of the forest. At night, they use tall dipterocarps on ridges for sleeping trees. The home range of a group studied in northern Siberut was 125 ha. Groups of up to 15 individuals have been observed in central Siberut, while larger groups of up to 27 individuals have seen in the north of the island. Groups are made up of one adult male, one or more adult females, and offspring. Siberut Macaques produce a diverse array of vocalizations throughout the day, including a loud, harsh bark produced by adult males, and given frequently around dawn. This call may function to indicate the direction of foraging in the morning. The Siberut Macaque occasionally interacts with sympatric langurs, sometimes forming mixed species associations with the Siberut Langur (Presbytis siberu). The most recent estimate suggests a density of ¢. 16 ind / km ® in the Peleonan forest of northern Siberut.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC9FFCCFFD5624FF654F9FF.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Vulnerable on The [UCN Red List. Legal protection of the Siberut Macaque in Indonesia is questionable because it is a newly recognized species. In recent years, forest clearing, logging, conversion to oil palm plantations, and the harvesting of non-timber forest products by local people have decimated its habitat. Hunting has increased with a burgeoning human population and improved access to remote areas due to logging roads and tracks. Rifles have replaced bows and arrows, and local rituals and taboos that formerly regulated hunting have been lost. Although not a preferred food, the Siberut Macaque is still hunted and poisoned because it is a crop pest. The pet trade is also a threat, and Siberut Macaques are kept and sold as pets in the Mentawai Islands. It occurs in one protected area, Siberut National Park, and is also abundant in Peleonan forest in northern Siberut, which is currently protected in the short-term by local agreements to exclude the area from logging concessions. Mentawai macaques in captivity in Indonesia may actually be Siberut Macaques rather than Pagai Macaques, but otherwise the Siberut Macaque has not been recorded in captivity. The total population is ¢. 17,000 - 30,000 individuals, down from ¢. 39,000 in 1980. An action plan published in 2006 suggested the following conservation actions: increased protection for Siberut National Park, which currently lacks enforcement; formal protection of the Peleonan forest in northern Siberut, which is home to unusually high primate populations and is easily accessible; conservation education, especially regarding hunting; and the development of alternative economic models for the local people to reduce the likelihood of selling their lands to logging companies.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC9FFCDFADC6357F997F5A9.taxon	materials_examined	Sumatra.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC9FFCDFADC6357F997F5A9.taxon	discussion	M. nemestrina has a characteristic helmet-shaped and bluntly bilobed glans penis, with the breadth of the glans 60 - 89 % of its length, that has resulted in assignment to various species groups of macaques, including the silenus-sylvanus species group (M. sylvanus, M. sinica, M. nemestrina, and the Sulawesi species) and more recently the nemestrina species group (M. nemestrina, M. silenus, M. leonina, M. pagensis). M. leonina and M. pagensis were formerly considered subspecies M. nemestrina. Distributions of M. nemestrina and M. leonina are contiguous in peninsular Thailand (between 8 ° N and 9 ° N) at the southern end of the Isthmus of Kra, and apparently restricted hybridization has occurred between them, including on the islands of Phuket and Yao Yai. Mid-Pleistocene fossils referable to the genus Macaca have been collected in eastern Java and may be ancestral to living M. nemestrina, an interpretation that implies that the pig-tailed stock inhabited Java during the Pleistocene and subsequently became locally extinct, as did other mammals, including the primates Siamang (Symphalangus syndactylus) and orangutan (Pongo). Prehistoric Holocene subfossils, collected on the Malay Peninsula and Sumatra, dating within the past 10,000 years, suggest a greater abundance of M. nemestrina in the western part of the Sunda area than presently occurs there. The Sunda Shelf (Sundaland) was most recently exposed during the last glacial maximum c. 18,000 years ago. Morphological similarity characterizes now disjunct insular and peninsular populations. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC9FFCDFADC6357F997F5A9.taxon	distribution	Distribution. Sunda area from the Surat Thani-Krabi depression in peninsular Thailand (8 - 9? N) SE through Peninsular Malaysia, Sumatra, Bangka, and Borneo; apparently native to offshore islets of Pinang (W coast of Peninsular Malaysia), Tioman (E coast of Peninsular Malaysia), and Batam (Riau Archipelago off the S tip of Peninsular Malaysia). It is thought to have been introduced to other offshore islets.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC9FFCDFADC6357F997F5A9.taxon	description	Descriptive notes. Head-body 53.2 - 73.8 cm (males) and 43.4 - 57.6 cm (females), tail 16 - 24 cm (males) and 13 - 25 cm (females); weight 10 - 13.6 kg (males) and 5.4 - 7.6 kg (females). The Sunda Pig-tailed Macaque is sexually dimorphic, males being a little less than twice the weight of females. Pelage of the trunk is yellowish-brown agouti to golden-brown agouti, including lateral surfaces of trunk and limbs and dorsal surface of appendages. There is a variably developed blackish mid-dorsal streak or patch that becomes more or less indistinct in the scapular region. Dorsal hairs are longest (7 - 9 cm) at the scapula in adult males. Indistinct dark streaks frequently occur on posterior surfaces of shanks. Underparts are thinly haired, whitish to ocherous-buff anteriorly, often becoming darker (buffy to pale brown) on the abdomen. Crown hairs are short and blackish, and they radiate to form a whorl at the top of the head. Crown patch is broad in front and extendslaterally on the supraorbital region as far as outer angles of the eyes. Cheek hairs are relatively short, pale at the base, and blackish at the tips, forming a pair of dark streaks or sideburns. The Sunda Pig-tailed Macaque has a long muzzle of pale brownish skin, thinly covered with inconspicuous short whitish hairs. Tail is thin, sharply defined blackish dorsally and pale ochrerous-buft ventrally; elongated terminal hairs frequently form an inconspicuous tuft. Tail is carried arched rearward and tip directed downward. Neonates are black. At about three months old, infant pelage changes to transitional brown and then to a tan agouti similar to adults.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC9FFCDFADC6357F997F5A9.taxon	biology_ecology	Habitat. Highest densities are found in lowland and hill dipterocarp rainforests. Although Sunda Pig-tailed Macaques occur from sea level to 1900 m, they prefer the higher elevations and dry ground at the foot of hills and slopes. They are present in lowland and swamp forest in Peninsular Malaysia. They are not regular residents of swamp forest in Sumatra but may enter freshwater swamps during months of lowest rainfall when many tree species are flowering or fruiting. They frequently occur adjacent to agricultural land, including hillside farms, and on fringes of urban environments. They may travel together, sometimes simultaneously, with Long-tailed Macaques (M. fascicularis) in lowland primary and secondary forest and degraded habitats.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC9FFCDFADC6357F997F5A9.taxon	food_feeding	Food and Feeding. Sunda Pig-tailed Macaques are mainly frugivorous but include leaves and invertebrates in their diet. During an 18 month study in Peninsular Malaysia, the diet consisted of fruits (742 %), leaves and buds (11: 1 %), flowers (11 %), invertebrates (12: 2 %), and other items (1: 4 %). Swarming termites, grasshoppers and other insects, and spiders were consumed elsewhere in Peninsular Malaysia, in addition to fruits, seeds, young leaves, leaf stems, and fungi. They eat figs (Ficus, Moraceae), other fruits, leaves, and miscellaneous food items in North Sumatra. Crop raiding is characteristic of the Sunda Pig-tailed Macaque. During a 14 month survey in Sumatra, more than 20 % of encounters occurred when the macaques were raiding farms. They appear to raid fields on a circuit, depending on the location and availability of crops. They concentrate on large fruits and exploit food resources that their relatively large size and strength assist them in obtaining — e. g. oil palm fruit, mature corn ears, thick and spiny-skinned durian fruit, and tapioca (cassava) roots. Sunda Pig-tailed Macaques also raid papaya, which can be quickly picked and transported to the forest edge for uninterrupted consumption. A field may be raided daily until a favorite crop, such as corn, is completely destroyed. They raid with stealth, frequently surveying a field for some time before entering it, one individual at a ime and rarely as a group. A lookout, often a subadult male, may keep watch from a tree at the forest edge, making a warning bark when humans approach. Infants taken from the wild and raised by humans are trained at 2 - 5 years old to respond to verbal commands to choose and pick coconuts. An efficient picker can harvest 500 - 1000 coconuts / day from a palm plantation. In Peninsular Malaysia, Sunda Pig-tailed Macaques sometimes are seen, frequently in association with Long tailed Macaques, scavenging through roadside garbage or begging for food from passing motorists.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC9FFCDFADC6357F997F5A9.taxon	breeding	Breeding. Sunda Pig-tailed Macaques show no synchrony or seasonality in sexual cycling. Mating occurs throughout the year, although there may be a slight peak in January — May. Females have a characteristic cyclical circumanal sexual swelling. Swelling develops gradually, beginning during or immediately after the menstrual period and reaches maximal tumescence in c. 15 days (at which maximal sexual behavior occurs in females and males). At maximum size, the engorged pinkish, and hairless area extends from the base oftail to the ventral border of the ischial callosities (seating pads), which are partly buried in the swelling, and extends laterally over an area twice the breadth of both callosities. Thinly haired skin anterior to the vulva (mons pubis) is less conspicuously swollen. Swelling rapidly subsides following ovulation, usually within 1 - 2 days after maximum tumescence. The first swelling may occur at 2 - 5 years old in captivity. The male’s broad, helmet-shaped glans penis has been described as “ pale crimson. ” The length of the baculum (penis bone) averages about 16 - 1 - 21 - 1 mm and the shaft is nearly cylindrical. The female’s cervix and cervical colliculi correspondingly are moderately large. In captivity, female Sunda Pig-tailed Macaques are sexually mature at three years old and males at 4 - 5 years old. Females typically solicit sexual advances of males by presenting their rump after approaching males from behind and standing in front of them. Males may initiate copulation with females by showing the “ pucker ” or “ protruded lip face, ” which is unique to the Sunda Pig-tailed Macaque. Copulation usually consists of a series of non-ejaculatory mounts that precede the ejaculatory mount (multimount ejaculator): 7 - 11 mounts are the median per ejaculation. A single offspring is born after a gestation averaging 172 days. Captive birth weights are 442 - 3 - 474 - 4 ¢ for female infants and 476 - 6 - 515 - 7 g for male infants. Age of weaning averages close to eleven months in captivity. Interbirth interval in natural populations may be two years. There is no record of infanticide in the wild. Known maximum longevity in captivity is 37 - 6 years.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC9FFCDFADC6357F997F5A9.taxon	activity	Activity patterns. Sunda Pig-tailed Macaques appear to spend more time traveling, and on the ground, than foraging and feeding. They flee quietly on the ground when disturbed. During a 14 month survey in Sumatra, 36 contacts were made while the macaques were traveling (38: 9 %) or fleeing (25 %) compared with feeding (5 - 6 %) or sitting (11 - 1 %). In another study in Eastern Kalimantan, traveling or foraging on the ground accounted for 67 % of contacts. An adjusted daily activity budget of locomotion (61 %), resting (19 %), foraging (16 %), and social behavior (4 %) was recorded during an 18 month study in tropical rainforest in Peninsular Malaysia. In north Sumatran forests, Sunda Pig-tailed Macaques generally forage in small, widely dispersed subgroups that travel on the ground and seldom remain in the same place for long; they search for miscellaneous food items on the ground but more often feed in trees. Contact is maintained with a low vocalization audible at distances of 30 - 80 m. A common pattern appears to be sleeping in broad-crowned emergents at the fringe of primary forest and entering secondary or degraded forest to raid cropland. Sunda Pig-tailed Macaques may vocalize as they approach sleeping trees at dusk. At other times, they can be so silent that they are difficult to observe.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC9FFCDFADC6357F997F5A9.taxon	biology_ecology	Movements, Home range and Social organization. Sunda Pig-tailed Macaques are wide ranging, with home ranges of at least 100 - 300 ha, parts of which may be used infrequently. Considerable home range overlap was reported in western Sumatra. Daily movements are ¢. 600 m in north Sumatra. Groups are multimale-multifemale. The adult male-female sex ratio has been found to vary from 1: 8 in 13 groups of 15 - 55 individuals (average 24) to 1: 2 - 7 (including adult and subadult mature males) in three groups of 49 - 81 individuals. Females remain in natal groups, and males emigrate at 5 - 6 years old. Both solitary adult and subadult males have been observed. Dominance hierarchies exist in both sexes. Adult males are dominant over females and may attack them at feeding sites, although groups of females (perhaps kin) may attack lower ranking males. Censuses in west-central Sumatra in 1996 - 1999 yielded density estimates of 1 - 7 groups / km? in lowland forest, 1 - 5 groups / km? in hill dipterocarp forest, 0 - 7 groups / km * in montane forest, and 0 - 8 groups / km? in submontane forest. Average group size was 10 - 5 individuals (range 1 - 20) in hill dipterocarp forest, seven individuals (range 6 - 8) in montane forest, 9 - 5 individuals in submontane forest, and 8 - 5 individuals (range 1 - 13) in lowland forest.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC9FFCDFADC6357F997F5A9.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Vulnerable on The IUCN Red List. The Sunda Pig-tailed Macaque has become seriously threatened and its populations fragmented by human encroachment and habitat loss, from legal and illegal logging, traditional and modern monoculture crop plantations such as oil palm, land clearance for agriculture and new settlements / transmigration, forest fires and drought (as have occurred in Kalimantan), and hunting for the illegal pet trade. Lowland forest 1 s most seriously threatened by a variety of human activities. Tropical rainforest, especially in lowlands, has disappeared rapidly in Sumatra, with most land being converted to commercial timber concessions, cultivated lands, and human settlements. The spread of oil palm plantations is one of the greatest threats to forests in Indonesia and Malaysia. From 1967 to 2000, the area under cultivation in oil palm in Indonesia expanded from less than 2000 km ® to more than 30,000 km ®. Indonesian production of crude palm oil was expected to reach a new high of 22 - 4 million tons in 2010, up 1 - 7 million tons in 2009. Production of palm oil is a major industry in Peninsular Malaysia, where Sunda Pig-tailed Macaques experienced an estimated decline of 43 - 7 % from 80,000 individuals in 1957 to 45,000 in 1975. Sabah and Sarawak have become the two big Malaysian oil palm states. Ultimately, the great majority of Sabah’s forest may consist of logged dipterocarp forest. There are many instances of conflict between humans and Sunda Pig-tailed Macaques, which has caused them to be regarded as a crop pest by farmers. Sunda Pig-tailed Macaques are found throughout Sarawak, for example, and can survive in logged forest but are hunted intensely for food and sport, and especially as crop pests in many areas. Along with the Long-tailed Macaque, they account for more than 2 % of mammals hunted in Sarawak and appear to be held at low density because of hunting. The Sunda Pig-tailed Macaque is naturally rare, and its groups are highly mobile so local populations are vulnerable to extirpation by hunting around farms. Trading of Sunda Pig-tailed Macaques for export by quota still occurs in Indonesia, and Sumatra is the main source. They are used as models for biomedical studies, including HIV / Aids research. Lower densities of Sunda Pig-tailed Macaques were recorded in Sumatra during surveys between 1996 and 1999 than those observed in the 1970 s. From 1977 to 1979, Peninsular Malaysia was the principal supplier of Sunda Pig-tailed Macaques to the United States, the world’s largest user of primates, supplying ¢. 970 individuals; imports from Indonesia totaled 570 individuals between 1976 and 1980. During 1978 - 2011, for which a record of international trade is available, more than 11,000 Sunda Pig-tailed Macaques may have been exported, with the United States being the major recipient. Numbers exported from Malaysia in 1977 - 1984 were significantly less. A total ban on the export of primates went into effect in Malaysia on 15 June 1984.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC8FFCEFAC66F8DF7EBF275.taxon	materials_examined	“ Mountainous and rocky situation, ” Arakan District, south-eastern Burma.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC8FFCEFAC66F8DF7EBF275.taxon	discussion	Until recently, M. leonina was classified as a subspecies of M. nemestrina. It has a characteristic helmet-shaped and bluntly bilobed glans penis, with the breadth of the glans 59 - 89 % of its length that has resulted in its assignment to various species groups of macaques. It is currently aligned with the nemestrina species group, including M. nemestrina, M. leonina, M. silenus, and M. pagensis. Distributions of M. leonina and M. nemestrina are contiguous in peninsular Thailand (between 8 ° N and 9 ° N) at the southern end of the Isthmus of Kra, and apparently restricted hybridization has occurred between them. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC8FFCEFAC66F8DF7EBF275.taxon	distribution	Distribution. NE India (S of the Brahmaputra River in the states of Assam, E Arunachal Pradesh, Manipur, Meghalaya, Mizoram, Nagaland & Tripura), extending SE through E Bangladesh and Myanmar (including the Mergui Archipelago), S China (SW Yunnan Province), Thailand, Laos, Vietnam, and Cambodia; S to the Surat Thani / Krabi depression in peninsular Thailand (8 - 9 ° N). The lack of records in C and NE Myanmar between 20 ° N and 25 ° N suggests that this may be a natural gap in the distribution of the Northern Pig-tailed Macaque.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC8FFCEFAC66F8DF7EBF275.taxon	description	Descriptive notes. Head — body 50 - 59.5 cm (males) and 40 - 49 cm (females), tail 18 - 25 cm (males) and 16 - 20 cm (females); weight 6.2 - 9.1 kg (males) and 4.4 - 5.7 kg (females). Dorsal pelage of the trunk of the Northern Pig-tailed Macaque is yellowishbrown agouti to golden-brown agouti anteriorly, becoming slightly drabber posteriorly, with lateral surface of the trunk and limbs similar in color. Hairs on the scapular region of adult males are 7 - 10 cm long. Chest and anterior surfaces of shoulders are a pale ocherous-buff, with underparts thinly haired, pale yellowish-brown to whitish, especially in females. Fur of the short, thin tail is blackish dorsally and buffy ventrally, with the tail base set off by a pair of projecting tufts of whitish fur. Tail is not invariably carried arched over the back, with tip directed upward and forward as frequently described, but may drop backward. The crown is short-haired, with a dark-brown patch extending in front about as far as the middle of the eyes. Cheek ruff is long and pale ocherous-buff, tending to hide the ears when seen from the front. A reddish streak extends laterally from the outer corner of each eye to cheek whiskers in males. Skin around eyesis a sharply defined pale bluish white. Muzzle is relatively short, with pale brownish skin densely covered with short buffy hairs. Pelage of adult females is somewhat shorter, paler, and drabber than that of males. Neonates are black. Pelage of infants changes to transitional brown a few months after birth. Immature Northern Pigtailed Macaques are generally paler and drabber than adult females, and the agouti pattern is less developed.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC8FFCEFAC66F8DF7EBF275.taxon	biology_ecology	Habitat. Dense evergreen and semi-evergreen forest, both tropical and subtropical, and deciduous forest in plains, foothills, and hills at elevations below 50 m to more than 2000 m. The Northern Pig-tailed Macaque is found in lowland primary and secondary forest and open habitats such as pine and dry dipterocarp forest and coastal, swamp, and montane forest. It is frequently observed in riparian habitat in western Thailand and Laos. It occasionally enters tea plantations in north-eastern India and has been seen in the vicinity of human settlements and cultivated fields in India and Laos, where it was relatively common in some areas in the 1990 s, even in highly degraded regions. The Northern Pig-tailed Macaque is similar in some morphological characteristics, behaviors, and ecological requirements to the Assamese Macaque (M. assamensis); their distributions overlap along the foothills of the South-east Asian mountain chains. The home ranges of two groups of Northern Pig-tailed Macaques overlapped with three groups of Assamese Macaques in Assam (India). Northern Pigtailed Macaques, specifically an adult male and female with an infant on her ventrum, have been observed traveling with a group of Assamese Macaques in a mosaic of deciduous and dry evergreen forest in western Thailand. The two species share habitat in Laos from 15 ° N to 20 ° N.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC8FFCEFAC66F8DF7EBF275.taxon	food_feeding	Food and Feeding. In eastern Assam, India, fruit comprises 65 - 9 % of the diet of the Northern Pig-tailed Macaque. They eat parts of more than 91 kinds of plants, including mature and tender leaves, leaf buds, leaf petioles, seeds, stems, climbers, roots, flowers, flower buds, inflorescences, bamboo shoots, and gums. They also eat insects and larvae, termite eggs, and spiders. Although crop-raiding is considered to be rare in north-eastern India, Northern Pig-tailed Macaques have been observed feeding in rice paddy after harvest as well as in fields of corn. They also eat fruits and vegetables grown by hill tribes in slash-and-burn cultivation (“ jhum ”). Recently, Northern Pigtailed Macaques have begun to beg for food, as they do in Khao Yai National Park, Thailand, where a few habituated groups may depend substantially upon biscuits, fruit, and other items from tourists.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC8FFCEFAC66F8DF7EBF275.taxon	breeding	Breeding. Northern Pig-tailed Macaques appear to have a marked winter-spring breeding season in north-eastern India. Females are sexually receptive in August-December, but a few females may be receptive until February. Newborns appear in mid-January to early May, with a few in June. Limited data suggest two birth peaks in western Thailand, approximately in June and December. Permanent canines erupt and puberty begins in males at c. 4 - 5 years old. Subsequently, the sizes of the glans penis and baculum (penis bone) increase rapidly, at a rate aboutfive times that of head-body length. The broad, helmetshaped glans penis is pinkish. The length of the baculum averages about 21 - 26 - 7 mm and is 25 % greater than in the Sunda Pig-tailed Macaque (M. nemestrina). The female's cervix and cervical colliculi correspondingly are moderately large. The sexual skin swelling is especially developed in female pig-tailed macaques and corresponds to their ovulatory cycle — reddish swelling occurs at mid-cycle and extends to the tail root. Swellings of Northern Pig-tailed Macaques resemble that of the Liontailed Macaques (M. silenus). They do not form a continuous pillow-like mass as in Sunda Pig-tailed Macaques but are subdivided into five separate swollen areas, with the subcaudal swelling being highly developed. A female's first swelling probably occurs at 2 - 5 - 3 - 8 years of age, as inferred from Sunda Pig-tailed Macaques and Lion-tailed Macaques. Females typically solicit sexual advances of males, presenting their rump after approaching males from behind and standing in front of them. Copulation usually consists of a series of non-ejaculatory mounts that precede the ejaculatory mount (multi-mount ejaculators). In a wild population, each mounting bout was 2 - 16 seconds, and number of thrusts was 3 - 23. A dominant male completed seven copulation bouts in two hours and 23 minutes but also 16 copulation bouts in 30 minutes. One female participated in copulation bouts for 20 days. After copulation bouts, the male sometimes utters a low bark. Post-copulatory grooming may occur; usually the female grooms the male. A single young is born after a probable gestation of 171 - 180 days. Gestation periods of 162 - 186 days and 167 - 179 days have been recorded in captivity. Captive individuals may live into their 30 s.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC8FFCEFAC66F8DF7EBF275.taxon	activity	Activity patterns. Northern Pig-tailed Macaques are primarily arboreal but may use both terrestrial and arboreal pathways, vocalizing to maintain auditory contact while traveling in a single file or in subgroups. They flee from danger through the trees, not on the ground. They do travel on the ground when crossing clearings and roads and to forage and feed in degraded areas and on crops. The amount of time spent on the ground to feed on crops fluctuates seasonally and by group. At one studysite in Assam, 38 % of the group’s time was spent on the ground in January when foraging in harvested rice paddy. At another site, as much as 20 % of daytime was spent on the ground in February-March feeding on sugar cane left over by foraging wild Asian Elephants (Elaphas maximus). Some detailed observations were made on the activity of Northern Pig-tailed Macaques in eastern and central Assam in 1992 - 1994 and 2004. From just after dawn, they had three peaks of feeding and a long midday rest, with shorter spells of resting in mid-morning and late afternoon. Average percentage of time spent in different activities was: resting (including preparation for night roosting) ¢. 45 %, feeding 23 - 5 %, locomotion 17 - 19 - 4 %, and grooming nearly 6 - 8 %. Actual time spent feeding was 158 - 229 minutes / day. Foraging for grain or seeds in harvested paddy occupied 65 % of feeding time in January in one area. In the presence of sexually receptive females, copulation became a major activity. A dominant male and a receptive female, who also copulated with a second male, spent as much as 33 - 7 % and 56 - 5 % of their respective time in copulation and related activities such as post-copulatory grooming and resting between copulatory bouts.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC8FFCEFAC66F8DF7EBF275.taxon	biology_ecology	Movements, Home range and Social organization. In eastern and central Assam, home ranges of Northern Pig-tailed Macaques are 83 - 347 ha, and overlap by about 25 - 48 %. Daily movement is 690 - 2240 m and is apparently influenced by weather conditions and the availability of seasonal fruit. Daily movement increased in degraded areas where fruiting trees were far apart. Groups are multimale-multifemale. The sex ratio recorded in Assam is 1: 5 - 5. It is assumed that males leave their natal groups at or before puberty to disperse to other groups. Both subadult and adult solitary males have been recorded. A linear dominance hierarchy among males was observed during the mating cycle in Assam, where the alpha male may lead group movement. Group size of the Northern Pig-tailed Macaque is 12 - 40 individuals. Exact counts of 16 - 33 individuals were recorded for seven groups in Assam. Estimated group sizes in Laos rarely exceed 20 - 30 individuals. There are a few records of groups numbering 50 - 150 individuals in Khao Yai National Park in Thailand, Dak Lak Province in Vietnam, and Dong Phou Vieng National Biodiversity Conservation Area in Laos. Such large groupings may be of two or more groups foraging and feeding together.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFC8FFCEFAC66F8DF7EBF275.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Vulnerable on The [UCN Red List. In South Asia, the Northern Pig-tailed Macaque is restricted to numerous fragmented locations, and appears to be few in numbers. There is little left ofits preferred habitat of dense forest. It is considered critically endangered in Bangladesh and endangered in India. Forest destruction and degradation are caused by tree felling, human encroachment, monoculture tree plantations, fisheries (in Bangladesh), and slash-and-burn shifting cultivation by cultural minorities or hill tribes (jhum in India). The Northern Pig-tailed Macaque is killed for food by many hill tribes in north-eastern India that increasingly have access to automatic firearms. Poaching has been rampant in some areas such as the Assam-Mizoram border. In China, the Northern Pig-tailed Macaque is variously classified as rare and endangered and was thought to number c. 1000 about the year 2000. It was classified as vulnerable in the Red Data Book of Vietnam in the 1990 s. Poaching for traditional medicine, stimulated by increasing affluence in the country (and overall in East Asia), is now putting extreme pressure on all primate populations in Vietnam, and no new conservation measures have been introduced. Direct threats to the habitat include logging, gold mining, and shifting cultivation by minority peoples. Elsewhere in the Indochinese Peninsula, development of the economy and infrastructure are resulting in habitat loss and degradation. In Laos, the Northern Pig-tailed Macaqueis listed as potentially at risk. Hunting for food by villagers does occur, and it may be affected by opportunistic hunting to supply bones to Vietnam for traditional medicine. Although these threats are in incipient stages, agriculture (especially commodity crops), mining, and hydropower are causing significant loss of forest habitat in Laos. In Cambodia, where Northern Pig-tailed Macaques are reported in only five of 24 provinces, use in traditional medicine, loss of habitat from logging, and especially trade constitute major threats to primate populations. Economic land concessions and mining concessions occur even within protected areas. Logging, plantations, aquaculture, hunting for food and pets, and political unrest in cultural minority states impact primates, including Northern Pig-tailed Macaques, in Myanmar. Factors associated with substantial economic development, such as timber extraction, large-scale farms and plantations, irrigation and hydroelectric projects, highway construction, mineral exploration, resettlement programs for hill tribes and others, and recreation and tourism have significantly reduced forest cover in Thailand. Shifting cultivation of hill tribes and ethnic Thais also has resulted in large areas of forest being cleared. Illegal hunting for sale (market hunting) and poaching for food near villages can directly impact primate populations, including Northern Pigtailed Macaques. The Western Conservation Corridor in western Thailand may offer some protection to both habitat and wildlife.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFCAFFCFFFF1640AF8A6F9D2.taxon	materials_examined	no type locality.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFCAFFCFFFF1640AF8A6F9D2.taxon	discussion	M. sinica is a member of the sinica species group of macaques, including M. radiata, M. assamensis, M. munzala, and M. thibetana. Albinism has been reported. There are some intergradations between subspecies. Three subspecies recognized.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFCAFFCFFFF1640AF8A6F9D2.taxon	distribution	Subspecies and Distribution. M. s. sinicaLinnaeus, 1771 — NE & SESriLanka (DryZone). M. s. aurifronsPocock, 1931 — SWSriLanka (WetZonelowlandsfromthecoastto ¢. 2000 mabovesealevel). M. s. opisthomelas Hill, 1942 — SC Sri Lanka (montane areas above elevations of 1800 m).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFCAFFCFFFF1640AF8A6F9D2.taxon	description	Descriptive notes. Head-body 44 - 53 cm (males) and 40 - 45 cm (females), tail 51 - 62 cm (males) and 46 - 57 cm (females); weight 4.1 - 8.4 kg (males) and 2.3 - 4.3 kg (females). The Toque Macaque is the smallest of the macaque species. It is yellow-brown or golden brown on the upperside and has pale undersides. The head has a whorl of dark elongated, radiating crown hairs that form a central cap extending to the frontal brows and to the nape behind. Face is pink in males and red in females; ears, eyelid borders, and lower lips are painted black in both sexes. It has a relatively long, slender tail that is 100 - 144 % ofits head-body length and black or dark gray above but paler underneath. Age-specific morphological changes are well pronounced in the Toque Macaque. Subspecific differences in morphology occur; e. g. the cap of “ Dry Zone Toque Macaque ” (M. s. sinica) is golden brown, but the “ Wet Zone Toque Macaque ” (M. s. aurifrons) has strong yellow anterior crown hairs that contrast with a darker brown posterior. The “ Highland Toque Macaque ” (M. s. opisthomelas) is the largest of the subspecies, with relatively short and stout limbs.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFCAFFCFFFF1640AF8A6F9D2.taxon	biology_ecology	Habitat. A variety of forest types such as gallery and semi-deciduous forests up to elevations of 2100 m. Toque Macaques are generally found in areas around permanent water. The Dry Zone Toque Macaque is found in dry evergreen forests, the Wet Zone Toque Macaque in lowland and midland tropical rainforest and wet-zone lowland forests (up to 3000 mm rainfall / year), and the Highland Toque Macaque in montane tropical rainforests. The Toque Macaque occasionally lives in human settlements, and it is protected around Hindu and Buddhist temples.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFCAFFCFFFF1640AF8A6F9D2.taxon	food_feeding	Food and Feeding. The Toque Macaque is largely frugivorous, and its diet is complemented with seeds, leaves, flowers, and animals such as insects, lizards, birds, and small mammals. Groups are known to raid garbage dumps, crops, and granaries.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFCAFFCFFFF1640AF8A6F9D2.taxon	breeding	Breeding. Female Toque Macaques have a 28 day reproductive cycle. The perineum reddens without swelling, and females secrete a pungent mucus from their vaginas, which signals that they are receptive. Polyandry is common, and malesinitiate copulation. Birth usually takes place at night in the canopy but occasionally on the ground during morning hours. A single offspring is born after a gestation of ¢. 168 days. Offspring have soft black natal coats and undeveloped crown caps, and they are hairless on their chests and abdomen. Nursing of the young usually lasts twelve months. Placentophagy is observed; the birthing mother, alpha female, and other female members of the group eat the placenta. Males reach sexual maturity at 5 - 7 years of age, and females reach sexual maturity at 4 - 5 - 6 years. Toque Macaques can live up to 30 years.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFCAFFCFFFF1640AF8A6F9D2.taxon	activity	Activity patterns. Toque Macaques are largely arboreal. They sleep in forks of branches in tall trees away from the trunk.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFCAFFCFFFF1640AF8A6F9D2.taxon	biology_ecology	Movements, Home range and Social organization. Toque Macaques live in multimale — multifemale groupsof ¢. 20 individuals, and occasionally up to 30 - 40 individuals. Their social organization is similar to that of the Lion-tailed Macaque (M. silenus), the Bonnet Macaque (M. radiata), the Stump-tailed Macaque (M. arctoides), and the Barbary Macaque (M. sylvanus) in that it is relatively peaceful and conciliatory (higher rates of tension-reducing contacts) compared with the strong nepotism and strong linear hierarchies of the Rhesus Macaque (M. mulatta) and the Japanese Macaque (M. fuscata). The extent of kin-bias in social relationships is also lower; females are more inclined to contact, groom, and form coalitions with unrelated females. The “ silent bared-teeth display, ” used to demonstrate submission and subordination in Rhesus and Japanese macaques, is used by Toque Macaques as an apparently friendly gesture and as a sign of submission. Male Toque Macaques are, however, more aggressive than females, and they dominate them across all age classes. Alpha males are known to usurp and dominate resources over subadult males and low-ranking females. Such aggression and competition often result in male emigration and the formation of new groups. The highest-ranking female in the group enjoys the highest reproductive sex, and she maintains a close relationship with breeding males during periovulatory periods and at other times. Post-reproductive females are withdrawn from most social activity, but they contribute by leading the group to unfamiliar ground and also in locating lost members of the group. Juveniles associate more with breeding females than postreproductive females. Group fission is common, with large groups breaking up into smaller groups. Following such fissions, the reproductive success of individual females increases, with formerly low-ranking females getting better access to food. They also experience decreased competition and reduced aggression by adult males and highranking females. Low-ranking females form peripheral groups that are later joined by subadult males or alpha males that have lost their standing due to aggression and competition in their former groups. These peripheral groups eventually move away to form groups of their own in separate home ranges. Male emigration is common and is usually a result of intragroup competition for mates and resources. Females are philopatric and continue living in their natal groups as they age. Subadult and juvenile females display “ aunting ” behavior (handling and taking care of their infant sisters), and unlike Rhesus and Japanese macaques, they may also take infants of unrelated breeding females. As in other macaque societies, even adult males are attracted to infants.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFCAFFCFFFF1640AF8A6F9D2.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Endangered on The IUCN Red List, including all three subspecies. There is no legal protection for the Toque Macaque under Sri Lankan law. All three subspecies are threatened with widespread habitat loss and by capture for the pet trade; they are even used as practice targets by the Sri Lankan and Tamil military. Relentless habitat loss within their distribution has caused a decline of ¢. 50 % of the overall population — a trend thatis likely to continue due to continuous loss of forest cover. The population is also affected by other stochastic events. Dry Zone Toque Macaques occur in numerous protected areas, including Flood Plains, Kaudulla, Maduru Oya, Minneriya, Yala, Somawathiya, Udawalawe, Wasgamuwa, and Wilpattu national parks. They also occurs in various sanctuaries, including Buddhangala, Kanthale Naval Sanctuary, Polonnaruwa, Randenigala, Ritigala Strict Nature Reserve, Sirigiriya, and Victoria-Randenigala-Rantambe Sanctuary, as well as Menikdena Archaeological Reserve, Elehara Forest Reserve, and Remmalakanda Forest Reserve. In south-western Sri Lanka, the hill country suffered an 80 % loss ofits forests in the late 19 " century, with their transformation into tea plantations and agricultural expansion for other crops. Wet Zone Toque Macaques have smaller group sizes and larger home ranges than the other subspecies, and as a result they are suffering increasingly from conflict with humans. They are found in Bellanwila-Attidiya, Muthurajawela, and Kurulu Kelle sanctuaries; Dombagaskande, Kitulgala-Kelani Valley, Rammal a Kanda, and Sinharaja forest reserves; and Thangamalai Wildlife Reserve. Highland Toque Macaques of central Sri Lanka are restricted to ¢. 400 km? of remaining habitat but occupy only ¢. 100 km? *. Ongoing habitat loss (coffee and tea plantations and pastures) is a serious threat to the Highland Toque Macaque.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFCAFFC8FAFF6369F89CF582.taxon	materials_examined	India.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFCAFFC8FAFF6369F89CF582.taxon	discussion	M. radiata is a member of the sinica species group of macaques, including M. sinica, M. assamensis, M. thibetana and M. arctoides. It is sympatric with M. silenus in the western part of its distribution. Albinism has been reported. Two subspecies recognized.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFCAFFC8FAFF6369F89CF582.taxon	distribution	Subspecies and Distribution. M. r. radiataE. GeoffroySaint-Hilaire, 1812 — S & WIndia (statesofGujarat, Maharashtra, AndhraPradesh, Goa, Karnataka, Kerala, andTamilNadu), theNlimitistheTaptiRiver, StothePalniHillsandSEasfarasTimbala, inlandofPuducherry (= Pondicherry). IthasbeenintroducedtotheMascareneIs, includingMauritiusandRéunion, probablyinthe 16 " century. M. r. diluta Pocock, 1931 — SE India (states of Kerala & Tamil Nadu), from the S tip and the SE coast, N to Kambam at the SW foot of the Palni Hills and Puducherry (= Pondicherry) in the E coast.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFCAFFC8FAFF6369F89CF582.taxon	description	Descriptive notes. Head-body 51.5 - 60 cm (males) and 34.5 - 52.5 cm (females), tail 51 - 69 cm (males) and 48 - 63.5 cm (females); weight 5.4 - 11.6 kg (males) and 2.9 - 5.5 kg (females). Head-body length of adult males average c. 15 % greater than those of adult females, and weights of adult males average c. 75 % greater than those of adult females. The Bonnet Macaqueis a lightly built, rather long-tailed species. The coat is gray-brown to golden-agouti-brown above (becoming duller posteriorly), with slightly paler limbs and a darker dorsal surface of the tail. Underside is buffy or whitish and sparsely haired, with skin showing through. The head has a hatlike crown of hairs radiating from a central whorl to form a circular cap, with posterior hairs longer than anterior hairs and with short forehead hairs parted along the midline. This “ bonnet ” effect is further accentuated by a high, somewhat balding forehead. Face and ears are bare, with skin of adults being brown to pinkish or bright scarlet in some females. Tail is 88 - 136 % of head-body length. The “ Dark-bellied Bonnet Macaque ” (M. r. radiata) is dull gray-brown above with yellow tones, a dark median stripe posteriorly, and ocher-gray limbs. Its crown is grayish brown, with hairs on the cap often black-tipped. Ventral skin is dark bluish-gray. In the “ Light-bellied Bonnet Macaque ” (M. r. diluta), the upper bodyis bright yellow or golden-brown (slightly duller posteriorly), with pale gray-brown limbs. Crown is pale yellowish-brown. Ventral skin is whitish and largely lacks pigment, hence the name light-bellied macaque.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFCAFFC8FAFF6369F89CF582.taxon	biology_ecology	Habitat. Predominantly in lowland, wet evergreen, dry deciduous forests and bamboo up to an elevation of 2100 m and usually near forest edges and along rivers. Bonnet Macaques are commensal with humans and forest dwelling. They have a high tolerance to anthropogenic disturbance, flourishing in urban areas, railway yards, temple compounds, villages, alongside roads (where there are trees nearby), plantations, and farmland. In larger numbers in urban areas, they are considered pests. They are able to live in cultivated areasif they have a few large trees to live in, most notably favoring fig trees such as the banyan tree (Ficus benghalensis, Moraceae) and F benjamina. While Dark-bellied Bonnet Macaques are wide ranging and a commensal, the Light-bellied Bonnet Macaque has a restricted distribution, is forest dwelling, and not as abundant. They spend c. 30 % of their time on the ground as commensals with humans, and only c. 10 % in their natural habitats, the tall forests.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFCAFFC8FAFF6369F89CF582.taxon	food_feeding	Food and Feeding. The diet of the Bonnet Macaques is eclectic and includesripe fruits, seeds, flowers, shoots, young and mature leaves, pith, and animals (insects, spiders, frogs and lizards, and bird eggs). As is typical of macaques, Bonnet Macaquesfill their cheek pouches with food to chew and swallow it later. Fruits are a ready source of energy, and they are known to feed on mangoes, lantana, and wild figs. They also extract other nutritional supplements from other plant parts such as calcium from mature leaves and protein from young leaves. Preferred flowers are from Cullenia exarillata (Malvaceae), Knema attenuata (Myristicaceae), Fahrenheitia zeylanica (Euphorbiaceae), Loranthus (Loranthaceae), and Acacia auriculiformis (Fabaceae). Bonnet Macaques are also known to feed on leaves and buds of Xylopia parvifolia (Annonaceae), Mesua ferrea (Calophyllaceae), Diospyros buxifolia (Ebeneceae), and Ochlandra (Poaceae). Their diet consists of 70 - 1 % fruits, 12 -: 8 % flowers, 10 - 6 % leaves, 4 - 2 % leaf buds, and 2 - 2 % seeds. To compensate for the lack of protein in their predominantly fibrous diet, they feed on insects. On occasion, they raid crops such as peanuts, potatoes, beans, carrots, radishes, squash, rice, peas, grains, cauliflowers, coffee, and coconuts. In urban areas, where they generally experience food shortage, they raid garbage dumps and home gardens.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFCAFFC8FAFF6369F89CF582.taxon	breeding	Breeding. The menstrual cycle of the Bonnet Macaque is ¢. 28 days (mean 27 - 8 days), and it is marked by a reddening of the perineum. Bonnet Macaques are strongly seasonal breeders, with most mating occurring in July-September and most births in February-April. Subordinate males are not excluded from mating, and there is no evident consort behavior, but dominant males copulate most and will sometimes attack subordinate males caught in the act. Malesinitiate and solicit mating, and pairs leave the group temporarily to mate, as far as 500 m away. Pheromonal cues indicate when a female is receptive. Females terminate copulation as and when they want. Groups with just one male can be susceptible to what is termed a “ male influx, ” during their brief mating season. A recorded case involved a group of five females with just one crippled male. Three receptive females approached and mated with out-group males. These males invaded the group and forcibly mated with two females that had suckling young from the previous year, injuring the females in the process and resulting in death of their infants (at least one of the infants was killed by one of the males). At this time, the resident male left the group, returning only when all but two of the outgroup males had left. Two females and two males took up residence as a result ofthis influx. This event was not seen as typical or characteristic of the Bonnet Macaque, but the result of occasional opportunism as a breeding strategy by males. The gestation period lasts 162 days. Births occur during the dry season in January — May, with a peak in February-April. Single offspring are born to females about every year. Neonates are generally almost black. Young are nursed for 8 - 12 months, and sexual maturity of both males and females occurs at 2 - 5 - 3 - 5 years old. The female first gives birth at c. 4 - 5 years old. Mean interbirth interval with a surviving infant is 458 - 6 days, but when an infant is lost, it drops to 382 - 5 days. Despite reaching sexual maturity, males are not socially mature for another 2 - 3 years. Lactating females have a dark red face compared with non-lactating females that have a pale pink face. Individuals may live for ¢. 30 years.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFCAFFC8FAFF6369F89CF582.taxon	activity	Activity patterns. The Bonnet Macaque is diurnal, arboreal, and terrestrial. Daytime activity in natural forested areas is ¢. 90 % arboreal and ¢. 10 % terrestrial. In more open vegetation, such as cultivated areas, it is more terrestrial; about 30 % ofits daytime activity. At night, they sleep in trees, except in urban areas where they sleep on roofs and ledges. A year-long study ofa forest group provided the following activity budget: 37 % foraging, 30 % engaged in social activities, 21 % resting, 7 % traveling, and 5 % engaged in miscellaneousactivities. Feeding is particularly intense in the early morning and late afternoon. Much oftheir terrestrial foraging is for animal prey. Urban groups suffering food shortage and being exposed to predators and, at least in many cases, harassment from people spend more time on the move (20 %) than rural and forest groups.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFCAFFC8FAFF6369F89CF582.taxon	biology_ecology	Movements, Home range and Social organization. Group sizes of Bonnet Macaques in forests and farmland and along roadsides average 29 individuals (range 7 - 75). Depending on the region and customs (extent of persecution or provisioning), groups in urban areas are a little smaller and average 21 individuals (range 10 - 43), or they may be considerably larger, averaging more than 40 individuals. Groups are multimale — multifemale. Groups have 1 - 8 adult and subadult males and 2 - 11 adult and subadult females, with a mean sex ratio of reproductive adult males and females of 1: 1 - 2. Home ranges of different groups overlap with others (5 - 25 % but sometimes 100 %), and encounters between groups are frequent. Males are generally quite tolerant; they often huddle together, have a high frequency of affiliative social interactions with each other, and havelittle difficulty immigrating among groups. The hierarchy among males is weak and not very linear. Bonnet Macaques have frequent intergroup encounters, and adult and subadult males collaborate in defending their females and their territory (food resources). Males disperse, and females are generally philopatric, although females will change groups in the wild. Solitary individuals have not been observed. All-male groups are infrequent, but they have been seen (e. g. Darkbellied Bonnet Macaque in Bandipur and Mudumalai national parks). Home ranges average ¢. 200 ha in natural areas and c. 100 ha in urban areas. In the wild, male Bonnet Macaques lead the group, control fighting between group members, give alarm calls (vigilance), and defend the group in times of danger. Males form coalitions, targeting individuals with which they had had a previous conflict, but otherwise male — male relations are generally placid. Resting clumped in passive contact is typical of the Bonnet Macaque, in contrast with Sunda Pig-tailed Macaques (M. nemestrina) and Lion-Tailed Macaques (M. silenus) that huddle infrequently. Male — female mounting is solicited by the male, and the male’s position in mounting is seen as a display of dominance. Female — female mounting and male-male mounting have also been observed. Alloparenting is common, with females showing interest in and trying to carry and care for infants of other females. Natural predators are believed to include Tigers (Panthera tigris), Leopards (P. pardus), small cats, large raptors, crocodiles, large snakes, and, particularly in urban areas, domestic dogs. Group members cooperate when fighting predators. Bonnet Macaques have been observed chasing Golden Jackals (Canis aureus).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFCAFFC8FAFF6369F89CF582.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Least Concern on The [UCN Red Last, including both subspecies. Bonnet Macaques are hunted in parts of their distribution, and live individuals are occasionally traded for use in biomedical research and road shows. They are viewed as pests by farmers and horticulturists in somevillages and semi-urban areas, resulting in conflicts. Crop-raiding Bonnet Macaques are shot or captured and transported to distant forests. The southward distributional extension of the Rhesus Macaque (M. mulatta) poses some threat to the smaller Bonnet Macaque through competition and aggression. Although they are listed as Least Concern, the Bonnet Macaque has undergone significant declines throughoutits distribution. One long-term study indicated a decline of 41 - 5 % in the last 20 years and another study in commensal areas, a decline of 21 - 5 % in 20 years. Although Bonnet Macaques appear to adapt well to urban environments and temple grounds, a survey of 107 temples and tourist spots in the state of Karnataka found macaque groups at 37 of them and found they had been eliminated from 51, most particularly in the coastal region. There is a high probability of further declines in conflict zones in the nearto mid-term future. Further research and surveys are needed to reassess the status of Bonnet Macaques in southern India. The Dark-bellied Bonnet Macaque is known to occur in numerous national parks, including Bandipur, Bannerghatta, Eravikulam, Kudremukh, Mollem, Mudumalai, Mukurthi, Nagarhole, Sanjay Gandhi, Silent Valley, Sri Venkateswara, and Nagarjunasagar Sirisailam, and a large number of wildlife sanctuaries. The Light-bellied Bonnet Macaque occurs in Periyar National Park and nine wildlife sanctuaries: Grizzled Squirrel, Kalakkadu, Mundanthurai, Neyyar, Peechi Vazhani, Peppara, Periyar, Point Calimere, and Shendurney.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFCFFFCBFF2E6408FC2BF821.taxon	materials_examined	India, Assam.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFCFFFCBFF2E6408FC2BF821.taxon	discussion	M. assamensis is a member of the sinica species group of macaques that also includes M. sinica, M. radiata, and M. thibetana; their distributions are allopatric or parapatric and extend from Sri Lanka to east-central China. The glans penis of this group is highly distinctive: the apex is sagittate, it is narrowed towards the tip, and its length is less than 30 mm. The subspecies recognized here are clearly different in the relative tail length of the adult males. A population differing in general body form and coloration has been identified in Nepal and may represent a new subspecies. Two subspecies recognized.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFCFFFCBFF2E6408FC2BF821.taxon	distribution	Subspecies and Distribution. M. a. assamensisMcClelland, 1839 — S & SEAsia, 200 - 2750 mabovesealevel, EofthegreatbendoftheBrahmaputraRiver, inSWChina (SEXizangAutonomousRegion [= Tibet], SWYunnan, Guizhou, SWGuangxiprovinces), NEIndia (EArunachalPradesh, EAssam, Nagaland, Meghalaya, M. a., Mizoram, andTripurastates), SandEthroughN & EMyanmar, N & WThailand, Laos, andNVietnam. M. a. pelops Hodgson, 1840 — Himalayas up to 3100 m above sea level, from C Nepal (W limit Tipling, 83 ° 36 ’ E) E through NE India (N West Bengal, Sikkim, W Assam states), and Bhutan (E limit M. a. River, 90 ° 58 ” E), with a widely disjunct record, of what may be a geographic relict, in coastal SW Bangladesh (Sundarbans).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFCFFFCBFF2E6408FC2BF821.taxon	description	Descriptive notes. Head-body 53.2 - 73 cm (males) and 43 - 7 - 58: 7 cm (females), tail 19 - 36 cm (males) and 17 - 29.3 cm (females); weight 7.9 - 16.5 kg (males) and 4.9 - 8.7 kg (females). The Assamese Macaque is much larger than the southern species of the sinica species group and more sexually dimorphic. Sexual dimorphism of headbody length of male “ Eastern Assamese Macaques ” (M. a. assamensis) may exceed that of the “ Western Assamese Macaque ” (M. a. pelops). Pelage on dorsal and lateral surfaces of the trunk is brown, varying individually from golden brown to dark chocolatebrown, usually with scapular regions somewhat brighter, paler, and longer-haired than lower back regions. Bright burnt-orange coloration (erythrism) may occur on the anterior dorsal trunk during the cool season in western Thailand. Short, coarse, drab dorsal pelage is reported late in the dry season. Color of outer surfaces of limbs and tail is similar to the adjacent surface of the trunk. Ventral surfaces of trunk and limbs are thinly haired, pale buffy to whitish, with exposed ventral skin pigmented pale bluish. Crown is about the color of the back, with hair variably smooth, tufted, or forming a rudimentary cap. Buffy to whitish cheek tufts and chin whiskers usually prominent in adults, with blackish marginal hairs around the face. Face is thinly haired, with pale pinkish to whitish skin above the orbits and brownish to purplish skin on the muzzle. Early infant pelage is distinctively drab grayish brown.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFCFFFCBFF2E6408FC2BF821.taxon	biology_ecology	Habitat. Subtropical mid-elevation montane broadleaf evergreen rainforest. In Bhutan, the preferred habitat of the Assamese Macaque is mostly tropical wet evergreen and semi-evergreen forest in foothills and subtropical and temperate broadleaf forests in higher hills and mountains. In western Thailand, it has also been observed in mixed deciduous / bamboo and dry evergreen forest, patchily distributed from lowland to mountainous areas, and in hill forest. Limestone outcroppings, rocky cliffs, or protruding crags with sparse vegetation, sometimes near watercourses, are commonly important features of various habitats and may be preferred sleeping sites of the Assamese Macaque. In hill forest in rolling plateau (1000 m above sea level) in western Thailand, protruding rocky crags, or fig trees (Ficus, Moraceae) adjacent to them, may be preferred sleeping sites. Two large accumulations of macaque feces were recorded near cliff faces suggesting habitual use as sleeping sites. Frequently, the Assamese Macaque is, or was, the most common macaque in hills and mountains throughout its distribution. Human encroachment is causing a change in the landscapes where it is found. In West Bengal, some groups are semi-wild or provisioned, inhabiting areas adjacent to or in human settlements. At a dam site in Kanchanaburi Province, western Thailand, near its known southernmost distribution (14 ° 41 ° N), Assamese Macaques resort to begging and nuisance behavior to obtain food from tourists.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFCFFFCBFF2E6408FC2BF821.taxon	food_feeding	Food and Feeding. Assamese Macaques are predominantly vegetarian, although animal food, including insects and small vertebrates such as lizards, may be consumed. Fruit appears to be the principal natural food. Bamboo shoots are also eaten. Diets of mostly leaves are reported in Bangladesh and Nepal. Crop raiding, especially for corn, occurs in many regions throughout the distribution of Assamese Macaques. Figs are among fruits consumed in western Thailand. When feeding in trees, they gather food either by breaking off terminal branches that bear several fruits or by plucking individual fruits. They feed on fruits of the same species as the Lar Gibbon (Hylobates lar). Human disturbance of their native habitat is resulting in the use of new foods. In Darjeeling District, West Bengal, the Assamese Macaque eats fruits, leaves, seeds, petioles, bases of leaves, flowers, rhizomes, epiphytic roots, and cotyledons from 63 different plant species, of which 52 are wild and the rest agricultural. Fruits such as banana, orange, apple, mango, guava, papaya, litchi, cucumber, and tomato and items such as bread, biscuits, snacks, rice grain, grams, groundnuts, and molasses may be obtained directly or indirectly from humans. One study group located away from human habitation near riverbank and thick vegetation consumed 90 - 1 % natural food daily. A second study group inhabiting open-valley land interspersed with human settlements and tea plantations consumed less (743 %). Artificial foodstuffs made up the remainder of the diet in both groups. A preference for natural vegetation may persist despite gradual dependence on artificial food items.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFCFFFCBFF2E6408FC2BF821.taxon	breeding	Breeding. Cyclical ovarian swelling of sexual skin in females of the sinica species group is relatively modest. In the Assamese Macaque, swellings are reported to occur only in adolescence, although a subcaudal swelling has been noted during pregnancy in captivity. Gray, blue, or purple coloration of the sexual skin has been noted in all four species in the sinica species group, and a bluish perineal streak has been reported in a pregnant captive Assamese Macaque. A small number of records available from eastern India and Nepal to northern Vietnam suggests that births probably peak in summer (rainy season), and thus mating probably peaks in winter (dry season). A distinct four-month mating season (October — January / February) occurs in wild populations in north-eastern Thailand. Based on fecal hormone analysis, it appears that female Assamese Macaques synchronize mating, not ovarian cycles; receptivity 1 s socially mediated (significantly associated with the number of other females mating on a given day) and induced before onset of ovulation and maintained after conception. Conceptions are spread over the mating season, with few occurring at the beginning. Females are promiscuous, initiating 70 % of copulations and rarely refusing to copulate. Males are single-mount ejaculators. Extended sexuality may enhance benefits that a female receives from many males, although she may mate repeatedly with a “ primary partner. ” Harassment by other females is rare. A single young is born every 1 - 2 years after a gestation of ¢. 5 - 5 months. A captive female conceived until c. 18 years of age, but her last viable infant was born when she was c. 14 years old. Longevity in captivity is ¢. 28 years.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFCFFFCBFF2E6408FC2BF821.taxon	activity	Activity patterns. Assamese Macaques are rarely observed on the ground in the forest but will raid cultivated crops and explore areas of human habitation (both rural and urban). When frightened or disturbed, they may flee through the canopy and then descend to the ground and withdraw quietly through underbrush, or flee first through trees and then on the ground. In Darjeeling District, the time spent in daily feeding was nearly 1 - 5 hours in two study groups, with 51 % of this time occurring during a morning feeding session. Maximum feeding occurred in trees at heights of 5 m and 13 m. Although it varies seasonally, overall feeding time averaged 13 - 3 % of the daily routine. An activity budget recorded in Nepal in March-April during two successive years averaged 45 - 5 % feeding, 27 % walking, 16 - 5 % resting, and 11 % grooming. A provisioned free-ranging group of Assamese Macaques in Golpara, India, spent 36 % of its time feeding, 30 % resting, 20 % grooming, and 15 % moving, and in a second study during the same year spent 43 % foraging, 22 % walking, 18 % resting, and 17 % grooming. An estimated time budget for a provisioned group at a temple in northern Thailand in the month of September was ¢. 31 % resting, 27 % traveling, 17 % feeding, 15 % playing, 8 % grooming, 1 % engaged in aggressive behaviors, and 0 - 3 % engaged in sexual activities. Differences in nutritive values of foods may be responsible for differences in percentages of time spent feeding and foraging.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFCFFFCBFF2E6408FC2BF821.taxon	biology_ecology	Movements, Home range and Social organization. Home ranges of all species in the sinica species group may be relatively stable. In some areas (Sikkim, Darjeeling, India, and Guangxi, Yunnan, China), Assamese Macaques migrate seasonally between higher elevations in summer and lower elevations in winter. Groups forage for seasonally available fruit in phalanx or file formation in a mosaic of mixed deciduous / bamboo and dry evergreen forest in western Thailand. Typical of the genus, Assamese Macaques live in multimale-multifemale groups. Male transfer is assumed. Group size averages. 20 individuals (range 7 - 50 individuals). Single individuals (probably males) have been seen in Bhutan and Nepal. The mean sex ratio of adult males to adult females was 1: 2 - 3 in a small sample of three groups in West Bengal and northern Thailand and 1: 1 - 6 and 1: 1 - 9 in seven groups in Nepal during successive years. Another group studied in northern Thailand consisted of 53 individuals, including 13 males and twelve adult females, and 55 individuals, including 15 males and 14 adult females, in consecutive years. Large adult males may be important in detecting and providing protection against potential predators, including humans.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFCFFFCBFF2E6408FC2BF821.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Near Threatened on The IUCN Red List, including both subspecies. The extent of human population pressure and economic development differentially affect the status of the Assamese Macaque throughoutits distribution. Reduction and degradation of habitat and hunting and poaching constitute the most serious threats. Human encroachment on its native habitat has increased crop raiding and other pest behaviors. The Assamese Macaque is the most abundant primate in Bhutan. Existing levels of harvesting of forest resources are unlikely to be of serious concern. There is no organized poaching or trade of any primate in Bhutan. Rapid development in some areas and hydroelectric projects and the presence of villages in protected areas may constitute problems in the future. In contrast, the Assamese Macaque in India and Bangladesh is considered endangered and critically endangered, respectively. During the past 20 years, populations there have declined significantly because of habitat fragmentation and degradation, along with hunting for food and traditional medicine. Selective logging is a major cause of habitat loss, and the loss offruiting trees and human encroachment contribute to the decline in habitat quality. In 2002, the number of adult males and females was estimated at less than 300. In China, the Assamese Macaque was quite numerous and common in south-eastern Yunnan in the 1950 s and 1960 s but subsequently became rare. A significant decline began in Guangxi in the 1980 s. Before declaration of the 1988 Wildlife Protection Law, in which the Assamese Macaque was listed as a Protected Animal of the First Class, its bones and fur were commonly brought by state purchasing stations for traditional medicine. Local hunters continue to poach Assamese Macaques as if they were the Rhesus Macaque (M. mulatta), which can be trapped and hunted legally as a crop raider. Body parts of the Assamese Macaque are not uncommon at county markets. Traditional shifting cultivation by cultural minorities has long been a major disturbance to wildlife habitat but on a minor scale compared with technologybased agriculture, including rubber plantations. In China, the Assamese Macaque is now confined mainly to the upper elevations of steep mountains and karst hills with relatively degraded vegetation. It is classified variously as endangered or extremely threatened in China and may have numbered no more than 5000 - 7000 individuals around the year 2000. In Laos, the Assamese Macaque is usually the most common macaque in hills and mountains and is considered to be potentially at risk. It may be affected by opportunistic hunting to supply bonesfor traditional medicine in Vietnam. Although such threats are incipient, agriculture, especially commodity crops, mining, and hydropower, are causing significant loss of forest. The Assamese Macaque was classified as vulnerable in the Red Data Book of Vietnam in the 1990 s. Poaching for traditional medicine, stimulated by increasing affluence in Vietnam and East Asia, is putting extreme pressure on all primate populations in Vietnam, and no new conservation measures have been introduced. Direct threats to the habitat include logging, gold mining, and shifting cultivation by minority peoples. The Assamese Macaque may be losing preferred habitat because limestone outcroppings are being mined for the production of cement for the construction boom in the country. Factors associated with substantial economic development, such as timber extraction, large-scale farms and plantations, irrigation and hydroelectric projects, highway construction, mining, resettlement programs for hill tribes and others, and recreation and tourism have significantly reduced forest cover in Thailand. The shifting cultivation of hill tribes and ethnic Thais has also resulted in large areas of forest being cleared. Illegal hunting for sale (market hunting) and poaching for food near villages can directly impact primate populations including Assamese Macaques. The Western Conservation Corridor in western Thailand may offer some protection to both habitat and wildlife. Forest exploitation, hunting, and political unrest of cultural minorities have impacted Assamese Macaques in Myanmar.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFCEFFCBFFE06D36F897F89B.taxon	materials_examined	Zemithang (27 ° 42 ’ N, 91 ° 43 ’ E, at 2180 m above sea level), Tawang district, Arunachal Pradesh, India.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFCEFFCBFFE06D36F897F89B.taxon	discussion	M. munzala 1 s a member of the sinica species group of macaques, including M. sinica, M. radiata, and M. thibetana. It is sympatric with M. assamensis pelops, and recent publications indicate that it may be a variation of that species. A revision of the taxonomy of M. assamensis will help resolve the status of M. munzala. Given the paucity of information and much-needed detailed studies, however, M. munzala is retained here as a distinct species. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFCEFFCBFFE06D36F897F89B.taxon	distribution	Distribution. NE India (W Arunachal Pradesh State), in the districts of Tawang and West Kameng at elevations of 2000 - 3000 m; it possibly extends into Bhutan and Xizang Autonomous Region (= Tibet) in S China. There are reports of its occurrence in Mouling National Park in the Upper Siang District of C Arunachal Pradesh, but they have yet to be confirmed.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFCEFFCBFFE06D36F897F89B.taxon	description	Descriptive notes. Head-body 57 - 5 cm, tail 26 - 4 cm; weight 14 kg. The Arunachal Macaque is characterized by a dark, chocolate-brown body, dark crown patch, and a whip-like short tail. It has a prognathous head (jaws projecting beyond the upper part of the face), with the upper part of the face broader than the muzzle. There is a dark patch of hair on the temple, extending from lateral corners of the eyes or cheeks to the ears (a species-specific trait). Front of the crown has a prominent pale-yellow patch, with a central group of dark hairs forming a whorl or an erect tuft. Coat color varies seasonally. Features said to distinguish it from the Assamese Macaque (M. assamensis) include their extremely dark color, hairy underside of the body, relatively stocky short tail, dark brown facial skin, distinctive facial marks on the temple and forehead, and lack of prominent chin or cheek whiskers. All Arunachal Macaques have the prominent dark patch on their crowns, while Assamese Macaques have smooth short hair and pale crowns. “ Western Assamese Macaques ” (M. a. pelops) have longer tails than Arunachal Macaques.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFCEFFCBFFE06D36F897F89B.taxon	biology_ecology	Habitat. Dense Bhutan fir (Abies densa, Pinaceae) montane forests at elevations of 2000 - 3000 m (unconfirmed reports extend their range to as high as 3500 m) and a range of habitats such as degraded broadleaved forests, degraded open scrub forests, agricultural areas, and oak and conifer forest. No other macaque in the Indian subcontinent occurs at these high elevations. Arunachal Macaques are quite tolerant of human-modified areas, sometimes ranging close to villages and fields.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFCEFFCBFFE06D36F897F89B.taxon	food_feeding	Food and Feeding. Diets of Arunachal Macaques vary by season. High-quality nutritious foods are rare or unavailable in winter, and they subsist on a low-energy, fibrous diet, spending ¢. 75 % of their feeding time on pith from such species as Erythrina (Fabaceae) and flowering plants such as Elaeagnus parvifolia (Elacagnaceae); they spend more than 5 % of their feeding eating on bark. In spring, they feed on young leaves. To survive winter, Arunachal Macaques subsist on a less diverse but high-fiber diet in the absence of high-energy food types, and they increase the time they spend feeding to up to 66 % of the day.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFCEFFCBFFE06D36F897F89B.taxon	breeding	Breeding. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFCEFFCBFFE06D36F897F89B.taxon	activity	Activity patterns. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFCEFFCBFFE06D36F897F89B.taxon	biology_ecology	Movements, Home range and Social organization. Social structure of the Arunachal Macaque is matrifocal, and relations among groups are generally tolerant and peaceful. The average group size is 21 individuals (range 5 - 43). Home ranges contract in winter, sometimes restricted to areas with Erythrina. Densities in the Zemithang area of the Tawang District are 0 - 94 groups / km? or 22 ind / km?.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFCEFFCBFFE06D36F897F89B.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Endangered on The [UCN Red List. The Arunachal Macaque has yet to be listed on the Indian Wildlife (Protection) Act of 1972. There are considerable differences in population estimates, with one source estimating more than 550 individuals and another estimating less than 250 mature individuals in less than 5000 km ®. Anthropogenic disturbances such as tree felling, livestock grazing, and the collection ofleaflitter are some of the habitat threats. A major threat comes from their persecution as crop raiders (e. g. of maize and millet). They are snared and shot because of this. In one instance alone, 54 Arunachal Macaques were killed in a village in West Kameng District. They are sometimes kept as pets. Culturally sensitive and participatory awareness programs, crop alternatives to avoid financial losses to villagers, mitigating retaliatory killing due to crop raiding using deterrents and better crop protection measures, crop compensation and insurance programs, and a thorough understanding of crop-raiding patterns could be appropriate crop-raiding mitigation strategies. Many of the people of Tawang and high elevations of West Kameng belong to Monpa communities that are predominantly Buddhist and do not hunt macaques for food. The Arunachal Macaque occurs in Tsangyang Gyatso Biological Reserve and probably Mouling National Park of north-eastern India.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFCEFFD4FAEB62BBF6D2FA8D.taxon	materials_examined	China, near Moupin (= Baoxing), Sichuan.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFCEFFD4FAEB62BBF6D2FA8D.taxon	discussion	M. thibetana is a member of the sinica (or sinica-arctoides) species group of macaques, including M. sinica, M. radiata, M. assamensis, and arguably M. arctoides. This species hybridizes with M. mulatta in the Kowloon Hills, Hong Kong. Studies of molecular and morphological differentiation among populations are ongoing and indicate the need for future subspecies designations. C. P. Groves, in 2005, listed the subspecies esau described in 1912 from the west of Lochangho, Guangdong Province, China, and two subspecies described in 1996 — huangshanensis from Mount Huangshan, southern Anhui Province, eastern China, and guiahouensis from Guizhou Province, south-west China. Considered monotypic here.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFCEFFD4FAEB62BBF6D2FA8D.taxon	distribution	Distribution. EC China (25 - 33 ° N, 102 ° 30 ° - 119 ° 30 " E) in E Xizang Autonomous Region (= Tibet), Sichuan, S Gansu, S Shaanxi, Hubei, Anhui, Zhejiang, N Yunnan, Guizhou, Jiangxi, Fujian, N Guangxi, and N Guangdong provinces; W limit in the Yangtze Gorge in W & NW Sichuan and S limit at 23 ° 48 ’ N, ¢. 110 ° E in Guangxi. The species may range into NE India (Arunachal Pradesh, Assam, and Meghalaya states), although these reports appear to be based on misidentifications.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFCEFFD4FAEB62BBF6D2FA8D.taxon	description	Descriptive notes. Head-body 61 - 71 cm (males) and 51 - 63 cm (females), tail 8 - 14 cm (males) and 4 - 8 cm (females); weight 14.2 - 18.3 kg (males) and 9 - 13 kg (females). Exceptionally large male Tibetan Macaques may exceed 30 kg. This is the largest of the macaque species. It has a short tail, long dense fur, and a hint of a shoulder mane. Body is dark brown above (weakly or not at all banded) and pale buffy or gray below, with the underside sparser than the upperside, but still not very sparse. Limbs become paler distally to pale brown on hands and feet. Crown also is a paler brown, with a small whorl on the vertex and short hairs radiating from it. There is a prominent pale buffy beard and full cheek whiskers. Facial skin is pale brownish on the muzzle, with narrow naked bands of whitish on either side of the nose. Tail is only 7 - 9 % (adult male) and 11 - 13 % (adult female) of head — body length. Male Tibetan Macaques have small testes in a white or creamy-white scrotum, and the penis is distinct in shape from that ofall other macaques. Body hair of both sexes darkens with age, as does the skin on the faces of the males. Females’ faces redden with age.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFCEFFD4FAEB62BBF6D2FA8D.taxon	biology_ecology	Habitat. Broadleaf, evergreen, primary, and secondary forests but also subtropical and deciduous forest formations; seasonal environments at elevations of 600 - 2500 m. Tibetan Macaques are tolerant of human disturbance. They have been studied in Baishuihe National Nature Reserve and Mount Emei (both in Sichuan) and Huangshan Range (Anhui). The latter two sites are tourist destinations, and Tibetan Macaques there are regularly provisioned, which facilitates research. H. Ogawa noted that while Tibetan and Rhesus (M. mulatta) macaques have overlapping distributions, the former species out competes the latter in high-elevation habitats, and the two species occupy different ecological niches.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFCEFFD4FAEB62BBF6D2FA8D.taxon	food_feeding	Food and Feeding. Tibetan Macaques eat mainly fruits, along with young and mature leaves, shoots, stems, stalks, roots, flowers, berries, seeds, exudates, mushrooms, eggs, and animal prey (including invertebrates, snakes, and birds). Their diets vary by season, with bamboo shoots and fruits being staples in autumn, insects are more important in late spring and summer, and mature leaves and bark during the remainder of the year. Q. K. Zhao reported that Tibetan Macaques were more folivorous than are other macaque species. Tibetan Macaques are sometimes fed by humans near temples and at tourist sites.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFCEFFD4FAEB62BBF6D2FA8D.taxon	breeding	Breeding. Female Tibetan Macaques have a 26 day reproductive cycle and show a slight reddening of the genitals during the periovulatory period, but they have no sexual swelling. There is a discrete birth season, with mating taking place in July-December. Temperature is positively correlated with the males’ fecal testosterone levels. Highand mid-ranked males copulate more than low-ranked males at this time of year. Tibetan Macaques mate throughout the year and engage in non-reproductive copulation, which may reduce conflict among males and increase the females’ access to preferred foods; males are more likely to feed with mating partners. During reproductive and non-reproductive sex, the male ejaculates following a single mount of the female. Typically, a single young is born after a gestation of 5 - 6 months. Twins are rare but have been documented at Huangshan Range. Infants are pale yellow, and their coat changes to tan by their second year. Adults of both sexes handle infants in a behavior called “ bridging ” in which infants are used by one adult monkey to initiate and / or maintain contact with another adult. Young are nursed for ¢. 12 months. Sexual maturity occurs at five (females) to seven (males) years. Alpha males are often young natal adults that prefer to mate with unrelated group members, but mother-son and sibling mating have been observed occasionally. Free-living individuals can live for more than 25 years. Available data on these parameters come from free-living but provisioned populations. Provisioning may affect several aspects of life history, including discreteness of mating and birth seasons, age at first reproduction, interbirth interval, and age at death.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFCEFFD4FAEB62BBF6D2FA8D.taxon	activity	Activity patterns. Tibetan Macaques are diurnal and mainly terrestrial. They prefer to sleep on rockycliff ledges or in trees. During snowy weather, they may sleep in caves. Huddling while sleeping increases with decreasing temperature.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFCEFFD4FAEB62BBF6D2FA8D.taxon	biology_ecology	Movements, Home range and Social organization. Tibetan Macaques live in multimale — multifemale groups of 15 - 50 individuals, in which adult females outnumber adult males. The number of males in each social group is notably high, with a sex ratio (males to females) of 1: 19. Female macaques are philopatric, while males disperse from their natal group. In some macaques, including Tibetan Macaques, males have secondary dispersal (s), usually into neighboring groups that include male kin. Neighboring macaque groups have overlapping home ranges. Macaque daily movements and home ranges vary widely by habitat type, resource distribution, and group size. With respect to the entire genus, B. Thierry reported day movements of 100 - 3000 m and home ranges of 0 - 12 - 22 km *. Despite uniformity across the genus in basic aspects of social organization, macaque species vary in qualities of aggression, extent of kinbased affiliation, occurrence of post-conflict reconciliation, maternalstyles, and reproductive behaviors within their social groups. Species can be positioned on a graded, social-style scale. Grade one characterizes species in which conflicts are unidirectional and follow a strongly defined dominance hierarchy and have high-intensity aggression, low rates of reconciliation after fights, strong kin bias, and protective maternal styles. Grade four species have the opposite qualities. Tibetan Macaques appear to be closer to the grade-one end of the spectrum in terms of rates and direction of aggression and lack of reconciliation, but they also exhibit qualities more typical of species classified in grade four, such as intense male — male affiliation, relaxed maternal style, and low rates of female kin bias. Tibetan and Barbary (M. sylvanus) macaques are famous for a particular affiliative behavior called “ bridging, ” in which an infant is used by one adult to initiate or maintain contact with another adult. Because groups of Tibetan Macaque include many males, they may use bridging and other behaviors (e. g. embracing and mounting) to maintain alliances, reduce aggression, and sustain their rank in the dominance hierarchy. Adult female Tibetan Macaques also use bridging to regulate their relationships.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFCEFFD4FAEB62BBF6D2FA8D.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Near Threatened on The IUCN Red List. The Tibetan Macaqueis listed in the Second Category of State Key Protected Wildlife List in the China Red Data Book of Endangered Animals. Habitat loss within the distribution of the Tibetan Macaque is now deemed stabilized. In areas where forest has been cleared for farmland, Tibetan Macaques sometimes raid crops and are hunted by local people. Populations are now distributed in 60 nature reserves where hunting is prohibited. Outside of reserves, targeted hunting is probably rare, but the occupation of remote mountainous regions by Tibetan Macaques makes regular monitoring difficult at some locations. Tibetan Macaques can adapt to various environmental conditions and can be found living in close association with humans or in human-disturbed environments. Most non-human predators (e. g. large cats) of the Tibetan Macaque have been extirpated. Tibetan Macaques are rare in captivity in and outside of China; they are difficult to maintain in a captive setting. No comprehensive field survey has been conducted, but the total population size has been estimated at ¢. 10,000 individuals. Tibetan Macaques are best known from research conducted at two tourist sites: Mount Emei and Huangshan Range. At Huangshan Range, recent research has focused on impacts of park management styles and provisioning on infant mortality, how exposure to tourists affects the macaques’ stress levels and rates of intragroup aggression, and potential for zoonotic disease transmission. Tibetan Macaques at Mount Emei compete for access to areas that overlap with tourist trails. More dominant groups monopolize trails with greaterlikelihood of pay off from tourist-provided food. Atthissite, they have become aggressive and can pose a danger to visitors. H. Ogawa noted that provisioning by Buddhist monks of Tibetan Macaques at Mount Emei may have occurred for several hundred years, making this the longest consistently provisioned primate population.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD1FFD5FA3660AAF930F8B9.taxon	materials_examined	Cochinchine, a region in southern Vietnam; based on the mounted skin of an adult male collected by P. M. Diard in June 1822, for which the exact place of collection is unknown.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD1FFD5FA3660AAF930F8B9.taxon	discussion	Morphological evidence (genital morphology, female sexual skin morphology, and facial skin color), copulatory behavior patterns, blood proteins, and distribution suggest that M. arctoides is derived from the sinica species group, more specifically from a M. thibetana-like ancestor. Some authors recognize subspecies based on pelage differences. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD1FFD5FA3660AAF930F8B9.taxon	distribution	Distribution. S & SE Asia, NE India S and E of the Brahmaputra River (Assam, Arunachal Pradesh, Meghalaya, Nagaland, Manipur, Mizoram, and Tripura states), SW China S of 25 ° N (Yunnan, Guizhou, Guangxi, and Guangdong provinces), N Myanmar, Bangladesh, Thailand, Laos, Cambodia, and N Peninsular Malaysia where it could be a recent colonizer dispersed from S Thailand as recently as 1959; it may be now extinct in Bangladesh where it was last recorded in 1982 and 1989. Known records are, at present, concentrated along N and S margins ofits distribution, and it is rare or absent in the C Indochinese Peninsula, where deciduous forests predominate. The species has been introduced into Hong Kong.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD1FFD5FA3660AAF930F8B9.taxon	description	Descriptive notes. Head-body 51: 7 - 65 cm (males) and 48.5 - 53 cm (females), tail 1.7 - 8 cm (males) and 1.7 - 6 cm (females); weight 9.9 - 15.5 kg (males) and 7.5 - 9.1 kg (females). The Stump-tailed Macaque is a moderately large macaque. Pelage is long and silky, with hairs in the scapular region of adult males up to 115 mm long. Dorsal pelage color usually is brown, occasionally reddish (erythristic) or blackish (melanistic); individually variable sporadic traits may occur in the same group. Outer surfaces of limbs are about the same color as the trunk, or slightly paler. Pelage of the shorttail varies from well-furred to partly hairless. Fur on underparts is thin and slightly paler than on the dorsal surface but considerably darker than the visible ventral skin. Crown hairs radiate from the center and are long at the back and sides. Anterior crown hairs are short and brown, creating a bald appearance. Side whiskers and beard are slightly paler than dorsal pelage. Face, including cheekbone regions, is thinly haired. Skin on orbital and cheekbone regions is pink to red, and areas around the nose and mouth are variably buffy to blackish. Adult females are about the same color as adult males. In old age, scattered gray hairs appear in both sexes, and facial skin may lose its reddish color. Stump-tailed Macaques are cranially distinguished by broad anteriorly directed cheekbones (malars) and relatively small canines. Newborn pelage is whitish, and facial skin is pale pink. In older infants, dorsal pelage begins to darken from whitish to buffy to brown on the lower back; dorsal surface of the trunk is brown by about one year of age. Infants also are distinguished by a characteristically strong odor.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD1FFD5FA3660AAF930F8B9.taxon	biology_ecology	Habitat. Primary broadleaf-evergreen rainforest in hilly areas and adjacent degraded and disturbed habitats. Stump-tailed Macaques can also be found in mixed deciduous / bamboo forest. The known elevational range is 50 - 2800 m, and there are no records from extreme lowlands. Climate throughout the distribution of the Stumptailed Macaque varies from tropical to subtropical. Chronically low food availability may characterize some of these habitats. Apparently it is totally absent from offshore islands; it 1 s not known to swim.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD1FFD5FA3660AAF930F8B9.taxon	food_feeding	Food and Feeding. Diets of wild Stump-tailed Macaques contain mostly vegetable materials, including fruit, seeds, flowers, leaves, and roots. They dig tubers and roots out of the ground. Two hundred plants were identified as food sources of Stump-tailed Macaques during a field study in Jorhat District, Assam, India; seeds and fruits constituted more than 45 % of the food intake. At least 18 species of cultivated plants are raided for food. Rice and especially corn were raided in an area of slash-and-burn agriculture formerly occupied by Hmong hill people at an elevation of ¢. 1000 m in lower montane forest in western Thailand. Stump-tailed Macaques opportunistically eat animal food, including insects (adults and larvae), freshwater crabs, frogs, birds, and bird eggs.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD1FFD5FA3660AAF930F8B9.taxon	breeding	Breeding. Reproductive anatomy of the Stump-tailed Macaqueis strikingly distinct in both sexes. The male’s glans penis is lanceolate and about twice as long as in other species of macaques, and the baculum (penis bone) is extremely elongated and straight or gently curved in form. Scrotum is red, and the penis is deep bluish-purple. The female’s prominent vaginal collicle, short vagina, and long exocervix (at least 50 % longer than in any other macaque species) are morphologically complementary to the genitalia of the male. The vulvar area of adult females likewise is red, and the clitoris is dark bluish-purple. The male’s testes descend into the scrotum at about three years of age, and the onset of semen production may occur about a year later. Onset of menstruation in captive females occurs about two years of age, first ovulation before age four, and first birth before age five. Sexual maturity in captive males and females, and presumably in natural populations, occurs 2 - 3 years before full adult size is reached. Captive females do not exhibit cyclical swelling of sexual skin or other obvious changes in perineal morphology during various phases of a 30 day ovarian cycle. Nevertheless, references to sexual skin swelling and receptive / proceptive behavior continue to occur in the literature. Copulatory behavior may be less dependent on female hormone levels than in other macaque species, but copulation frequency was observed to be variably elevated around mid-cycle in some colony groups. Copulation may be initiated by either sex. A female initiates copulation by presenting her hindquarters to a male. A male initiates copulation by approaching, touching, or threatening a female, thereby inducing her to present her hindquarters. A male responds to hindquarter presentation by visual, oral, olfactory, and digital examination of the female’s perineum, and subsequently mounts her dorsoventrally. Pelvic thrusting usually continues through a single uninterrupted mount until ejaculation occurs (single-mount ejaculation). In colony groups, numbers ofthrusts per ejaculation were 26 - 64 and took 36 - 59 seconds. A “ frowning round-mouthed expression ” is exhibited by males at ejaculation. Intromission usually continues while the male sits and pulls the female onto his lap (the unique “ pairsit ”), with mean duration of 67 - 114 seconds. Copulations in colony groups usually occur in short series that involve a single male-female pair and may be monopolized by competitive dominant males. Limited evidence suggests the possibility of seasonal breeding in natural populations. The mean gestation ofsingle young in 93 colony pregnancies was 177 - 3 days (extremes 161 - 210 days). Mean birth weight for infants in two colonies was 495 g. Infants in one laboratory colony were nursed for at least 7 - 9 months. Until a new infant is born, offspring may be accepted on the nipple even after lactation has ceased. The interbirth interval averages 1 - 6 years. In semi-natural groups, reproductive rate was estimated to be one infant per adult female every two years. Age of menopause is unknown, although reduction in reproductive capacity may occur before 17 years of age. Maximum known longevity in captivity is c. 30 years.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD1FFD5FA3660AAF930F8B9.taxon	activity	Activity patterns. Stump-tailed Macaques wake at about dawn; feed until 10: 00 - 11: 00 h; rest, groom, and play during midday; feed again in the late afternoon beginning ¢. 17: 00 h; and retire to one or a few sleeping trees. Percentages of time allocated to different activities in a group in Jorhat District were 39 - 9 % resting, 24 - 8 % feeding, 16 - 9 % traveling, 15 - 6 % grooming, and 2 - 4 % engaging in other activities. Stump-tailed Macaques usually forage and travel on the ground but are able to travel and forage in all levels of the forest canopy. Midday activities and feeding on seasonally available fruits, seeds, and leaves may take place in trees. In response to danger, they usually flee on the ground, but occasionally into the canopy. They can be aggressive in encounters with humans.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD1FFD5FA3660AAF930F8B9.taxon	biology_ecology	Movements, Home range and Social organization. Home ranges of Stump-tailed Macaquesin Jorhat District are 336 - 587 ha, with core areas of 193 - 320 ha. Daily movements of wild populations are 2 - 3 km in Yunnan, China, and 1 - 8 - 2 - 3 km in Assam, India; a semi-wild group in peninsular Thailand moved 0 - 4 - 3 km / day. Groups of Stump-tailed Macaques are multimale-multifemale. The ratio of adult males to adult females is c. 1: 6 in Tripura, India and 1: 1 - 4 in Assam, India. Twenty adult males and 16 adult females were recorded in a free-ranging group of 73 individuals in southern Thailand. Group sizes vary from 2 - 3 individuals to ¢. 60 - 70. A few solitary males have been reported in north-eastern India and western Thailand. It seemslikely that young males in wild populations leave their natal group at or before sexual maturity. There was one record of a group of 100 or more individuals in peninsular Thailand, and unverified verbal accounts of similar numbers elsewhere in peninsular Thailand and western Thailand in the 1970 s — - 1980 s.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD1FFD5FA3660AAF930F8B9.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Vulnerable on The IUCN Red List. The Stump-tailed Macaque appears to have been fairly common throughout much of its distribution as recently as the early 20 " century, but it has become generally rare because of the burgeoning human population and rapidly expanding economic development. In South Asia, it is considered critically endangered and may have become extinct in Bangladesh. The population in north-eastern India may have been reduced to seven subpopulations totaling less than 250 individuals because its specialized habitat has been degraded and fragmented through selective logging, timber and firewood collection for charcoal, deliberately set fires, and the construction of dams, roads, and powerlines, with consequent soil loss and erosion. Stumptailed Macaques are hunted for food and traditional medicine (their bones), and live animals may enter local pet trades. They are classified variously as endangered or extremely threatened in China, with a population numbering ¢. 3500 individuals. Overhunting, including exploitation for traditional food, has resulted in rapid shrinkage of its distribution in south-western China, and they may have been extirpated in Guangdong Province. Habitat loss from the exploitation of evergreen broadleaf forest is a serious threat. In the 1990 s, the Stump-tailed Macaque was considered to be widespread above 11 ° N in central Vietnam and was classified as vulnerable. Earlier, it had been selected to be used in the production of polio vaccine. Poaching for traditional medicine, stimulated by increasing affluence in Vietnam and the East Asian region, is now putting extreme pressure on all primate populations in Vietnam, and no new conservation measures have been introduced. Direct threats to the habitat include logging, gold mining, and shifting cultivation by minority peoples. Elsewhere in the Indochinese Peninsula, development of the economy and infrastructure is resulting in habitat loss and degradation. In Cambodia, where the Stump-tailed Macaque is found in only three of 24 provinces, economic land and mining concessions occur even in protected areas. In Laos, Stump-tailed Macaques are common at only a few sites within its countrywide distribution. Large groups may be affected by opportunistic hunting to supply bones to Vietnam for traditional medicine. Terrestriality makes Stump-tailed Macaques more vulnerable to snaring than other species of macaques. Although still incipient, agriculture (particularly commodity crops), mining, and hydropower development are causing significant loss of forest habitat. The trend toward large-scale farms and plantations and use of marine products has resulted in serious habitat loss and degradation in peninsular Thailand. In 1964 - 1976, the USA — the world’s largest user of primates — imported ¢. 30,000 primates from Thailand. Most were probably macaques, especially Stump-tailed Macaques, of which Thailand supplied more than 90 % of the 2883 individuals imported by the USA in 1971 and 1972. Most were used in biomedical and other research, but some entered the pet trade as “ miniature chimpanzees. ” The population in peninsular Thailand was significantly reduced by trapping for this trade. In 1975, Thailand imposed a ban on the commercial export of macaques (and other primates), effective in April 1976, which virtually eliminated Stump-tailed Macaques from international trade.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD0FFD6FAE9629DF8B9F5F8.taxon	materials_examined	Indonesia, Sumatra, (Bengkulu).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD0FFD6FAE9629DF8B9F5F8.taxon	discussion	The M. fascicularis species group of macaques, including M. fascicularis, M. mulatta, M. cyclopis, and M. fuscata, is differentiated primarily on the shape of the male genitalia. In M. fascicular, the apex of the glans penisis bluntly bilobed, and the breadth of the glans is 41 - 55 % of its length. The fascicularis species group has alternately been defined as including only M. fascicularis and M. arctoides. Ten subspecies are recognized based on complete (or nearly complete) discontinuity in variation of the same characters in the nominate species, M. f. fascicularis: color of dorsal pelage, crown, or outer thigh; lateral facial crest pattern; head-body length in adult males; and relative tail length of adult males. M. fascicularis is parapatric or marginally sympatric with M. mulatta for ¢. 2000 km from south-eastern Bangladesh, west-central Thailand, and southern Laos to central Vietnam (c. 15 - 20 ° N). Intermediate tail length and possibly noticeable geographic variation in dorsal pelage color, lateral facial crest pattern, head-body length, and skull length suggest hybridization in the latter three regions. All these characters in M. fascicularis north of the Isthmus of Kra (c. 10 ° N) tend to be intermediate between those in M. fascicularis south of the Isthmus and those in M. mulatta. Y-chromosomes of M. mulatta have spread at least 400 km south to c. 12 - 5 ° N within the distribution of M. fascicularis. The subspecies aureus has a contact zone with the nominate form fascicularis in the eastern and southern parts of the Isthmus of Kra and may intergrade also with the nominate form or M. mulatta in central and eastern parts of the Indochinese Peninsula. A contact zone between the subspecies philippinensis and fascicularis inhabited by mixed-phenotypic populations occurs in eastern and central Mindanao, southern Negros, and possibly nearby islands in the Philippines. Ten subspecies recognized.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD0FFD6FAE9629DF8B9F5F8.taxon	distribution	Subspecies and Distribution. M. f. fascicularis Raffles, 1821 — S Laos, S Vietnam, Cambodia, E & S Thailand (and offshore Is), S to the Malay Peninsula, Borneo, Sumatra, Java, Bali, and most but not all offshore Is, also extending into the Sulu Archipelago and Zamboanga Peninsula of W Mindanao, in the Philippines; probably artificially introduced in the Nusa Penida-Timor Is chain. M. f. atriceps Kloss, 1919 — Ko Khram I (= Khram Yai), off SE coast of Thailand. M. f. aureus E. Geoffroy Saint-Hilaire, 1831 — SW Bangladesh (Teknaaf Peninsula), S Myanmar (including the Mergui Archipelago), WC Thailand (S to ¢. 10 ° N), and Laos. M. f. condorensis Kloss, 1926 — SE Vietnam (Con Son and Hon Ba Is in the South China Sea). M. f. fuscus G. S. Miller, 1903 — Simeulue I, off NW Sumatra. M. f. karimondjawae Sody, 1949 — Karimunjawa I and presumably nearby Kemujan I, N ofJava. M. f. lasiae Lyon, 1916 — Lasia I, off NW Sumatra. M. f. philippinensis 1. Geoffroy Saint-Hilaire, 1843 — Philippine Archipelago N of ¢. 10 ° N. M. f. tua Kellogg, 1944 — Maratua I, off E Borneo. M. f. umbrosus G. S. Miller, 1902 — Nicobar Is (Katchall, Little Nicobar, and Great Nicobar Is).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD0FFD6FAE9629DF8B9F5F8.taxon	description	Descriptive notes. Head-body 37 - 63 cm (males) and 31.5 - 54.5 cm (females), tail 36 — 71 - 5 cm (males) and 31: 5 - 62.8 cm (females); weight 3.4 - 12 kg (males) and 2.4 - 5.4 kg (females). The Long-tailed Macaque is the smallest of the fascicularis species group. Adult males are generally larger than adult females; mean head-body length of males exceeds that of females by 13 %. Geographic variation in dorsal pelage coloration is noticeable, ranging from buffy to yellowish-gray to golden brown to reddish brown to dark brown to blackish. Erythrism of dorsal pelage varies in intensity in different regions. Crown is usually more brightly colored than the back. Crown hairs are directed backward and outward, sometimes forming a small, pointed central crest or, alternately, a darker cap. Outer surfaces of limbs fade from about the same color as the adjacent area of the trunk to pale grayish to pale golden brown at wrists and ankles. The dorsal surface ofthe tail is darker at the root (golden-brown or blackish) and gets progressively paler toward the tip. Pelage on under surfaces of trunk, limbs, and tail is thin and pale gray to whitish. Thinly haired facial skin is brownish to pinkish with sharply defined whitish upper eyelids (although hair may be concentrated on the upperlip or chin). Cheek whiskers usually sweep upward from near the angle of the jaw to the outside margin of the crown between the eyes and ears (transzygomatic crest pattern); less frequently, in some populations (or individuals), the crestis restricted to the posterior region of the lowerjaw (infrazygomatic cheek pattern). All dorsal pelage of newborn infants is conspicuously dark brown to blackish. Infant facial skin is bare, lacking pigment, and pinkish.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD0FFD6FAE9629DF8B9F5F8.taxon	biology_ecology	Habitat. Preferred habitats include seashore, along with mangrove forest, riverbanks and swamp forest, most commonly found at low elevations but up to ¢. 2000 m above sea level. A 1996 - 1999 survey in west-central Sumatra found Long-tailed Macaques only in lowland and hill dipterocarp forests at elevations of less than 800 m. They can live successfully in edge habitats and are often found near human settlements in fields, botanical gardens, plantations, temple precincts, fruit orchards, and vegetable gardens. Some populations may invade human habitation in search of food. Dependence on human sources of food is widespread in Thailand, and the only natural populations there may be confined to the Western Conservation Corridor, specifically Huai Kha Khaeng Wildlife Sanctuary in Uthaithani Province.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD0FFD6FAE9629DF8B9F5F8.taxon	food_feeding	Food and Feeding. Long-tailed Macaques are opportunistic feeders, omnivorous but primarily frugivorous. They consume leaves and other plant parts, invertebrates (crustaceans, bivalves, and snails near water and insects inland) and small vertebrates (fish). On the seashore, they select and transport rocks to crack open oysters to eat. Loss of forest habitat may result in crop raiding of rice and corn in rural areas and scavenging (and begging) for food in urban areas.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD0FFD6FAE9629DF8B9F5F8.taxon	breeding	Breeding. Seasonal peaks in copulations, pregnancies, and births occur among populations of Long-tailed Macaques, but timing varies geographically and between years at the same locality — apparently associated with fruiting cycles. Single birth occurs after a gestation of 160 - 168 days. Females probably become reproductively active at c. 3 - 5 years old, and their first infant may be born when they are c. 4 years old. Males usually begin to copulate as subadults at 5 - 6 years old, after they have left their natal troop. The mean duration of the menstrual cycle in 28 captive females was 30 - 9 days. Prominent swelling of the bare skin between the root ofthe tail and anus occurs at puberty, and skin around the vulva frequently reddens. In post-pubertal females, sexual swelling and reddening are generally less conspicuous and are highly variable in size and duration. In a well-studied population in northern Sumatra, mating and consortships of most females tended to increase with increasing intensity of one of these characteristics. Mating may be initiated by either sex by staring at the other and usually continue with the male’s examination of the perineum of the female. The male mounts the female by gripping her hips with his hands and her calves with his feet. Ejaculation may occur either as a result of a single mount or multiple mounts. During a 4 year period in northern Sumatra, overall annual birth rate (births / adult female) was 0 - 58. High and low birth rates tend to occur in alternate years in the wild, but captive females are capable of producing young every year. A non-provisioned female may produce c. 8 - 9 offspring during her reproductive life (age c. 4 - 20 years). In captivity, known maximum life span for the Long-tailed Macaque is 37 years. Mean birth weight in a large captive group was 318 - 2 g (+ SD 45 - 2 g) for 563 female infants and 347 - 5 g (+ SD 55 - 2 g) for 600 male infants. In captivity, nursing lasts 9 - 22 months, although 15 months appears to be average. Adult male infanticide may be one of the important causes of infant death. The gradual change from blackish neonatal pelage to adult coloration begins at 2 - 3 months of age when infants begin to obtain some of their food independently; the transition appears to be complete before the eruption of permanent first molars at c. 15 months old.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD0FFD6FAE9629DF8B9F5F8.taxon	activity	Activity patterns. The Long-tailed Macaque is diurnal. It often forages on the ground in coastal and riparian areas; 25 % of encounters in Sumatra and 26 - 5 % in East Kalimantan Province (Indonesian Borneo). Feeding occupies 13 - 55 % of daily walking hours. It is predominantly arboreal in inland forest, favoring the lower and middle forest strata; 98 % in Peninsular Malaysia and ¢. 97 % in East Kalimantan. Tall, relatively bare trees near a river may be preferred sleeping sites. Groups of Long-tailed Macaques may flee from danger on the ground or in the canopy or both. They are excellent swimmers and apparently enter the water for pleasure.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD0FFD6FAE9629DF8B9F5F8.taxon	biology_ecology	Movements, Home range and Social organization. Non-provisioned groups of Longtailed Macaques contain 5 - 100 individuals, although they may subdivide into smaller units to forage. Groups are multimale-multifemale, with resident lineages of female kin and males emigrating from their natal groups. Dominance / subordination hierarchies exist within and between lineages in females and among males. Known ratios of sexually mature males to sexually mature females are 1: 0 - 58 - 7 - 1. Predator detection may be a major determinant of social behavior and group size. Home ranges are quite variable at 12 - 5 ha to more than 300 ha. Home range overlap is generally slight and encounters between adjacent groups are relatively rare, although home range of a “ daughter group ” may overlap extensively with that of her natal group. Reported mean daily movements are 325 - 1900 m and may be positively correlated with group size.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD0FFD6FAE9629DF8B9F5F8.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Least Concern on The [UCN Red Lust. Subspecies condorensis and umbrosus are classified as Vulnerable, philippinensis as Near Threatened, fascicularis as Least Concern, and atriceps, aureus, fuscus, karimondjawae, lasiae, and tua as Data Deficient. Populations of Long-tailed Macaques have been decimated in Bangladesh by shrimp farming and shipbuilding, and habitat availability and quality have been significantly reduced in Myanmar. Throughout much ofits distribution, especially that of the nominate form fascicularis, human-macaque conflict is burgeoning in both rural and, more recently, urban landscapes as human populations expand and forest habitat is encroached upon. Responses have included culling (Peninsular Malaysia), increased quotas for trapping of wild monkeys for domestic research as well as “ breeding stock ” for primate export companies (Indonesia), and sterilization (Singapore) — frequently in the absence of surveys and population estimates, or using questionable survey data. At the same time, wild populations suspected of having been laundered as captive-bred may be exploited for the international trade driven by industrial testing and biomedical research. According to the CITES database maintained by the United Nations Environment Programme-Wildlife Conservation Monitoring Centre (UNEP-WCMC), more than 254,000 live Long-tailed Macaques were exported for medical, scientific, commercial, and breeding purposes from 2004 until July 2010. Imports of Long-tailed Macaques into the USA — the world’s largest user of primates and, as such, a bellwether of pressure on wild populations — increased from 17,124 individuals in 2004 to 24,629 individuals in 2005 and reached 26,799 individuals in 2008. A recent decline in numbers imported into the USA may be associated with outsourcing. Participants in the open meeting at the 2008 Congress of the International Primatological Society (IPS) in Edinburgh, UK, to review the biennial listing of the “ World’s 25 Most Endangered Primates, ” ajoint activity of the [IUCN / SSC Primate Specialist Group, Conservation International, and IPS, officially recognized the Long-tailed Macaque as the first “ widespread and rapidly declining ” primate species, and thelisting of the Long-tailed Macaque on The IUCN Red List is currently being reassessed.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD3FFD7FA2E6F5DF73DF274.taxon	materials_examined	India, based solely on Tawny Monkey, a menagerie animal (presumably observed in London), not preserved, identified by Pennant in 1771.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD3FFD7FA2E6F5DF73DF274.taxon	discussion	Restricted by R. Pocock in 1932 to “ Nepal Tarai ” (= Terai), the narrow plain that extends along the southern border of Nepal. M. mulatta is a member of the fascicularis species group of macaques, including M. fascicularis, M. cyclopis, and M. fuscata, exhibiting the characteristic helmetshaped and relatively long and narrow glans penis. Assignment to a separate mulatta species group of macaques with M. cyclopis and M. fuscata also has been proposed. As many as ten subspecies have been recognized in classifications published since 1932: mulatta, sanctijohannis, lasiotus, tcheliensis, vestitus, villosus, littoralis, brevicaudus, siamica, and memahoni. Differentiation of local and regional populations, based on descriptions of traditional characters such as overall size, tail length, pelage color and length, and molecular diversity is regarded as inadequate by some recent authorities to warrant formal recognition of subspecies. The interspecific boundary with M. fascicularis extends ¢. 2000 km across the Indochinese peninsula in the south-east. The two species share a considerable range between 17 ° N and 14 ° N in Vietnam and Laos (and probably Thailand). Hybridization has been inferred from morphological intermediates in absolute and relative tail length recorded in Thailand, Laos, and Vietnam and in some M. mulatta with noticeably less rufous hindquarters in southern Laos. Molecular studies differentiate three populations: Indian, Chinese, and Indochinese, the latter the apparent result of introgression from M. fascicular. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD3FFD7FA2E6F5DF73DF274.taxon	distribution	Distribution. S & SE Asia from ¢. 36 ° N (in Afghanistan, Pakistan, India, and China) S to ¢. 15 ° N (in India, Thailand, Laos, and Vietnam), also in Nepal, Bhutan, Bangladesh, Myanmar, and Hainan I. An isolated population at 40 ° 24 ’ N in NE China was extirpated in 1987.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD3FFD7FA2E6F5DF73DF274.taxon	description	Descriptive notes. Head-body 41 - 66 cm (males) and 37 - 58 cm (females), tail 12 - 5 — 31 cm (males) and 12.5 - 28 cm (females); weight 4 - 14.1 kg (males) and 3 - 10 kg (females). The Rhesus Macaque is an intermediate-sized member of the fascicularis species group. Head-body length and body weight tend to increase with latitude, while tail length generally decreases. High relative tail length (47 - 5 - 72 - 1 %) in some southeastern specimens has been interpreted as evidence of hybridization with Long-tailed Macaques (M. fascicularis). Pelage of the upper back of the Rhesus Macaque varies from yellowish gray to golden brown to burnt orange with crown, nape, and sides of the head about the same color as the adjacent upper back. Pelage of the lower back varies from golden brown to burnt orange to intense burnt orange. A black streak marks the front edge of the crown, and blackish hairs may fringe cheeks. Crown hairs usually are smoothly directed backward. Hairs on sides of the head usually form a small crest or whorl near the angle of the jaw (infrazygomatic crest). Thinly haired facial skin is pale brown to reddish, except for upper eyelids that lack pigment. Dorsal pelage on proximal parts of limbs is similar in color to that on the adjacent trunk; pelage color of limbs is less erythristic distally. Base of the tail is approximately the same color as the lower back and becomes dark brown dorsally and buffy ventrally. Ventral surfaces of the trunk and limbs are thinly haired and pale buffy to whitish. The Rhesus Macaque undergoes seasonal molting and fading of pelage coloration. Pelage in Indian populations at upper elevations (2400 m) may be longer and sleeker than nearby populations at lower elevations. Abnormally pale “ golden ” individuals reportedly occur in various parts of India and Pakistan (and perhaps Thailand). Newborns have pale brown to dark brown pelage, generally darker than in adults, and a midline bare area in the crown hair. Bare skin of the face, hands, and feet changes from dull purple to pale pink within about five minutes of birth. Skin color changes to buffy by about two months of age, and the dark neonatal pelage is gradually replaced by paler pelage similar to that of adults, and without the midline part, by about six months of age.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD3FFD7FA2E6F5DF73DF274.taxon	biology_ecology	Habitat. Deciduous and mixed deciduous / bamboo forest, scrub and secondary vegetation, and evergreen forest, in seasonal subtropical climates. The natural distribution of the Rhesus Macaque in the north extends to temperate and subalpine habitats, where forests are mixed broadleaf / needle-leaf and, less frequently, needle-leaf. It is highly adaptable and occurs in arid areas in western India and tidal swamps in eastern India and Bangladesh. Most elevation records are below 2000 m (below 600 m in Laos), but it occurs at ¢. 3200 m in Nepal and ¢. 4000 m in Qinghai Province, China. The Rhesus Macaque frequently inhabits disturbed areas, where it raids cultivated fields. In South Asia, especially India, it increasingly lives as a commensal with humans in urban and rural areas, and can be found at roadsides and temples.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD3FFD7FA2E6F5DF73DF274.taxon	food_feeding	Food and Feeding. Rhesus Macaques are primarily vegetarian, and their diets include fruits, seeds, flowers, leaves, buds, shoots, twigs, stems, roots, bark, pith, and gums from hundreds of species of angiosperms, gymnosperms, and fungi. Plant consumption varies geographically; as many as 150 wild plant species were recorded as food in northern India (Uttar Pradesh). Insects may be the most common animal food; other animal food includes spiders, crayfish, crabs, shellfish, fish, bird eggs, and honeycombs. Rhesus Macaques living in tidal swamp forest of the Sundarbans catch and eat fish on mudflats. Groups near human settlements opportunistically raid diverse cultivated crops and gardens and forage through garbage, while temple populations in India and Nepal may be provisioned.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD3FFD7FA2E6F5DF73DF274.taxon	breeding	Breeding. Reproductive behavior of the Rhesus Macaque is strongly seasonal in natural populations throughout its distribution, with mating peeking in autumn and winter and births occurring in spring and summer. Sexual maturity in females in natural populations usually does not occur until 3 - 5 - 5 - 5 years of age, although some may engage in fertile copulations as young as 2 - 5 years old. In captive females, first menstrual bleeding may occur about one year before sexual maturity. Males appear to become sexually mature somewhat later, usually at 6 - 5 years old. At ¢. 5 - 5 years old, the testes permanently descend into the male’s scrotum. At or before this age, males generally leave their natal group. In postjuvenile females and males, regions of the skin undergo intermittent swelling and reddening associated with the reproductive cycle. In captive females, a pair of pinkish pubic swellings develops during the second or third year of life. Following the first menstrual bleeding, a large bilobed, blister-like pubic swelling extending backward to the sides of the vulva marks the beginning of puberty. During adolescence, which may last up to two years, the sexual skin becomes less acutely swollen but redder and more extensive, often extending to the root ofthe tail and over the buttocks and posterior surfaces of the thighs and iliac region. Sexually mature females experience reddening rather than swelling of the sexual skin, which frequently includes the face and nipples. Maximum intensity of the red color generally occurs near the day of ovulation and is retained during pregnancy and frequently lactation. Red sexual skin develops in about the same posterior and facial regions in pubertal and adolescent malesas in females, but there is no pubertal swelling. The red color becomes brighter in males during the mating season. Female sexual activity is cyclical in natural populations. The periovulatory period is 8 - 12 days, and ovulation appears to occur at the midpoint. Females average more than two receptive periods per mating season, and post-conception receptive periods occur also. Temporary mating associations or consortships are formed between receptive females and male partners. The duration of consortships varies from ¢. 25 minutes to eleven days although 1 - 2 days may be typical, and females usually consort with more than one male. Consortships and mating may be initiated by either sex. A female typically presents her perineal region to the male who mounts her dorsoventrally. Usually, a series of mounts, separated by dismounts, is required to complete copulation. Coagulated semen or a “ vaginal plug ” may be visible on the female’s perineum following copulation. Male and female homosexual mounting and male masturbation have been observed in natural groups. The gestation period is 133 - 200 days and averages c. 166 days. A single infant is born. Birth weight in one large laboratory averaged 476 g in 1067 female infants and 502 - 8 g in 1115 male infants; birth weight in natural populations may be less. Mean annual birth rate in natural populationsis 42 -: 9 - 90 - 8 %. Infants may begin to take small amounts of solid food at two weeks of age. Mothers begin to resist nursing attempts about three months later, although nursing may extend through the subsequent mating and birth seasons and not be completed until the first infant is about one year old .. The greatest age at which a captive female is known to have produced a living infant is about 28 years and six months. Males may remain capable of copulatory ejaculation to c. 30 years old. Maximum reported longevity in captivity is 37 years.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD3FFD7FA2E6F5DF73DF274.taxon	activity	Activity patterns. Rhesus Macaques may spend as much as 72 % of daylight hours on the ground and 28 % in trees. Forest groups tend to be more arboreal than non-forest groups. Reportedly, they rarely descend from trees in the tidal swamp forest of the Sundarbans. In wild populations, average proportion of waking hours spent feeding is 15 - 8 - 45 %, and feeding generally peaks in the morning after waking and in the afternoon before roosting for the night.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD3FFD7FA2E6F5DF73DF274.taxon	biology_ecology	Movements, Home range and Social organization. Home range averaged c. 65 ha in 323 non-forest groups of Rhesus Macaques in rural and urban areas and 196 ha in 129 forest groups. Overlap of home ranges of adjacent groups is extensive, and most contact between groups seems to provoke some degree of antagonism. Different parts of a group’s home range may be used during different seasons, depending on the availability of food and water. Daily movements averaged 1 - 2 km in nine non-forest groups and 1 - 9 km in an estimated 16 forest groups. The mean size of c. 1182 non-provisioned or minimally provisioned groups was 32 - 2 individuals, with group sizes of two to ¢. 250 individuals. Non-provisioned groups tend be the largest (86 to ¢. 105 individuals) in the far north in Afghanistan and Qinghai and Henan provinces, China. Groups of Rhesus Macaques are multimale-multifemale, with resident lineages of female kin. Fission usually occurs between matrilines. Males usually leave their natal groups and join a nearby group at c. 5 years 6 months old at or before permanent testis descent into the scrotum; subsequent transfers may occur from a few months to a few years later. The sex ratio of adult males to adult females is c. 1: 3. Solitary males have been observed throughout the distribution of the Rhesus Macaque.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD3FFD7FA2E6F5DF73DF274.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Least Concern on The IUCN Red List. Wild populations of Rhesus Macaques were threatened by trapping for export in South Asia during the mid- 20 " century. During 1956 - 1960, the USA annually imported an average of 120,000 Rhesus Macaques from India, following a 1955 agreement between the two countries permitting their export for use in development and testing of polio vaccine and for other medical uses. An annual export quota of 50,000 individuals was subsequently established. In the state of Uttar Pradesh, the only area for which data on population trends are available, intensive trapping for export caused a conspicuous change in the age structure of the populations of Rhesus Macaques, with juveniles experiencing the greatest decline. Further reductions in the export quota were followed by an Indian ban on the export of all primates, effective in 1978, as the Indian governmentcited the USA for using Rhesus Macaques in militaryrelated research. During 1964 - 1978, the USA — the world’s largest user of primates — imported a total of 332,000 primates from India, of which more than 99 % probably were Rhesus Macaques. China began captive breeding of native Rhesus Macaques in 1978 and was exporting them by 1984. Molecular differences between Chinese and Indian macaques resulted in renewed interest by Western researchers in obtaining the latter. Efforts to establish breeding colonies in Nepal were abandoned because of local protest. Hunting, habitat loss, forestfires, accidental mortality, and changes in native species dynamics formerly posed threats. Subsistence hunting still occurs in northeastern and central India and mid-central Nepal, and there is some trade for pets and entertainment. Some trapping for a pet trade may also occur in eastern Afghanistan. Currently, the main threat is deterioration and loss of habitat and increasing conflict with people, especially in South Asia. The Indian government has invoked a policy of “ zero tolerance ” toward pest behavior in urban areas. In 2007, the northern Indian state of Himachal Pradesh embarked on a program of conflict management that includes masssterilization of Rhesus Macaques.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD5FFD0FF216405F973F585.taxon	materials_examined	Japan.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD5FFD0FF216405F973F585.taxon	discussion	M. fuscata is a member of the fascicularis species group of macaques, including M. fascicularis, M. mulatta, and M. cyclops, and has the characteristic helmet-shaped and relatively long and narrow glans penis. It would seem that M. fuscata arrived in Japan from a progenitor probably similar or identical to M. mulatta on the Korean peninsula to Kyushu-West Honshu during the Middle Pleistocene. There were evidently now-submerged land bridges during one or two glacial intervals of sea level depression ¢. 0 - 63 - 0 - 43 million years ago. The oldest macaque fossils found in Japan are a molar tooth and humerus similar to those of M. fuscata. Their age is difficult to ascertain, but through association with proboscidean fossils, they are tentatively dated to the age of one or the other of these land bridges. Korean macaque fossils are about the same age. Two subspecies recognized.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD5FFD0FF216405F973F585.taxon	distribution	Subspecies and Distribution. M. f. fuscataBlyth, 1875 — JapaneseArchipelagoSoftheTsugaruStrait (41 ° 30 ° N), inHonsu, Shikoku, andKyushuIs, andtheclose-lyingKojima (= Koshima), Kashima, Awajishima, Shodoshima, andKinkazanislets. M. f. yakui Kuroda, 1941 — Japanese Archipelago (Yakushima I, ¢. 60 km S of Kyushu).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD5FFD0FF216405F973F585.taxon	description	Descriptive notes. Head — body 47 - 2 — - 65 cm (males) and 46.4 - 60.1 cm (females), tail 8: 7 cm (males) and 8 - 1 cm (females); weight 5.6 - 18.4 kg (males) and 4 - 13.8 kg (females). Mean head-body length of male Japanese Macaques exceeds that of females by 9 %. Mean body weight of males exceeds that of females by 32 %. Color of the crown and relatively long dorsal pelage varies from pale yellowish brown to grayish brown to dark golden brown, with fur on outer surfaces of limbs and tail slightly paler than the adjacent trunk surface. Pelage varies from whitish to pale gray to pale ocherous buff on ventral surface of the trunk and inner surfaces of limbs. Prominent side whiskers are somewhat paler than the crown, and hairs on sides of the head usually form a small crest or whorl near the jaw angle (infrazygomatic crest). Thinly haired facial skin is pinkish to reddish. Tail is very short and well furred and may have a white tuft at the tip. Dorsal pelage color, generally slightly paler in females than in males, tends to be paler in the northern part of the distribution of Japanese Macaques, and an annual molt occurs in late spring — early summer. Dorsal pelage of newborn infants is dark brown to blackish, conspicuously darker than in adults, and pink facial skin is paler than in adults. Replacement of newborn pelage begins at c. 2 months of age and is complete by 4 - 5 months. The nominate subspecies has pale brown to dark brown pelage on dorsal surface of hands, similar to or paler than dorsal trunk pelage. The “ Yakushima Macaque ” (M. f. yakui) is distinguished by blackish pelage on the dorsal surface ofits hands, distinctly darker than its dorsal trunk pelage.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD5FFD0FF216405F973F585.taxon	biology_ecology	Habitat. Warm temperate, evergreen broadleaf forest in the southern part of the distribution and cool temperate, deciduous broadleaf forest in the northern part. The Japanese Macaque is absent in most lowland areas, although it is presumed to have inhabited these areas before intensive human exploitation. Elevational range extends to 3180 m. Observations have been made ofJapanese Macaques in subalpine needle-leaf forest at 3050 m during relatively mild weather. The so-called “ snow monkeys ” in the Shigakogen area of Nagano Prefecture survive winter snows by bathing in thermal pools.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD5FFD0FF216405F973F585.taxon	food_feeding	Food and Feeding. Japanese Macaques eat forest fruits, seeds, nuts, leaves, flowers, shoots, buds, and bark. Minor dietary components include roots, grasses, herbs, fungi, insects, crabs, spiders, mollusks, fish, and bird eggs. On Koshima Islet, they are known for the innovation and social transmission of food-preparation techniques, such as sweet-potato washing and wheat washing (“ placer-mining ”). Elsewhere, they are known for non-utilitarian behaviors such as stone handling among young individuals. Crop raiding by Japanese Macaques has intensified with human encroachment upon their habitat.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD5FFD0FF216405F973F585.taxon	breeding	Breeding. Male Japanese Macaques appear to sexually mature at c. 4 - 5 years of age, and by five years of age almost all males have emigrated from their natal group. Full socio-sexual maturity does not occur until ¢. 8 - 5 years when the frequency of mating activity is aboutfive times as great as at younger ages and the (near-) mature, intensely red sexual skin of the face, perianal area, and scrotum has developed. The first menstruation of females usually occurs at ¢. 3 - 5 years old, at about which time para-anal pubertal swellings and proceptive / receptive behavior may occur. Subcaudal or paravulval pubertal swellings occur during mating seasons over the next 2 - 3 years. Bare skin on the face, nipples, and perineum of adult females becomes intensely reddish during the mating season, increasing and decreasing in intensity during receptive and non-receptive intervals, respectively. An odoriferous vaginal discharge occurs during the periovulatory period in adult females, but there are no perineal swellings. A mature female may exhibit zero to six proceptive / receptive periods during the mating season. Conception usually occurs during the first ovulation cycle, but post-conception copulations may occur. The mean gestation is 171 - 7 days. A small sample of 21 birth weights suggests that mean weight in female neonates (546 - 8 g) is close to that in male neonates (538 - 7 g). In provisioned groups, females generally give birth to their first infant at ¢. 5 - 6 years. Mean interbirth intervals are 1 - 7 years in provisioned and 2 - 6 years in non-provisioned groups. Reproduction is strongly seasonal, with mating restricted to autumn-winter and births to spring — summer. Birth dates tend to occur earlier as latitude increases, presumably to enhance infant development before the poor feeding conditions in winter. During the mating season, females and males, including nongroup solitary males, form consortships of varying duration, and both sexes are likely to have more than one consortship partner during a mating season. A male usually mounts a female dorsoventrally, gripping her waist and shanks with his hands and feet. Ejaculatory copulations usually consist of a series of mounts separated by brief dismounts of ¢. 30 seconds (multiple-mount ejaculators). Non-reproductive sexual behavior (homosexual and autosexual) has been observed in natural and semi-natural groups. Infants may be fully weaned by 6 - 8 months or continue to nurse for about one year. The greatest known life span is 33 years in a provisioned group, and the greatest age at which a female is known to have given birth is 26 years.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD5FFD0FF216405F973F585.taxon	activity	Activity patterns. Japanese Macaques are semi-terrestrial. Detailed information on time spent on the ground and in trees is available from a study of the population at Tsubaki Wild Animal Park, Shirahama-cho, Wakayama Prefecture. During daytime hours, average time spent on the ground was 68: 3 % for males and 39 - 8 % for females. Feeding, including ingestion of mature broadleaf tree leaves after a season of unusually poor fruit production, occupied 38 % of daytime hours in a group in western Yakushima. The Japanese Macaque usually sleeps in trees, but sleeping on the ground has been reported on the two small islands of Kinkazan and Kojima, neither of which harbors predators. Japanese Macaques reportedly swim well, including in the ocean.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD5FFD0FF216405F973F585.taxon	biology_ecology	Movements, Home range and Social organization. Home range sizes of Japanese Macaques are strongly influenced by forest type and are positively correlated with group size. Home range is relatively small in evergreen broadleaf forest (1 - 4 - 6 - 4 ha / group member) and much larger in deciduous broadleaf forest (9 - 79 ha / group member). On western Yakushima, the population density was estimated at 0 - 7 groups / km? in undisturbed evergreen broadleaf forest and 1 - 5 groups / km? in disturbed evergreen broadleaf forest. The mean group size of 117 wild groups was 40 - 8 (+ SD 28 - 9) individuals (range 10 - 161). Provisioned groups tend to be larger; maximum group size reported in a provisioned group at Takasakiyama, Oita Prefecture, was 1255 individuals. Aggressive behavior between neighboring groups varies from frequent to rare, or absent. Groups ofJapanese Macaques are multimale — multifemale, with resident lineages of female kin and males more than five years old emigrating from natal groups and entering nearby or even distant groups. A daughter eventually outranks all females that rank below her mother. She remains subordinate to her mother, but newborn daughters rise in rank above their sisters’ rank. The mean composition of 35 natural groups was ¢. 18 % adult males, 32 % adult females, 35 % juveniles, and 15 % infants. Group size may change by fission or extinction by fusion. Large groups often split into two or three groups, reportedly along matrilines.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD5FFD0FF216405F973F585.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Least Concern on The [UCN Red List, including both subspecies. Prior to 1945, the range of the Japanese Macaque was contracting as an apparent consequence of habitat destruction and hunting for food and medicinal purposes. It disappeared from the small islands of Omishima, Hiradoshima, and Tanegashima in the mid- 20 " century. Subsequent population expansion has been evident in eastern Japan, while both expansion and reduction (or disappearance) of local populations have been observed in western Japan. The status of populations ofJapanese Macaques in less suitable habitat such as higher mountain slopes remains unknown. Hunting regulations have protected them from commercial and sport hunting since 1947, but the same law provides for removal of nuisance animals such as crop-raiders; the number of which has risen rapidly since the early 1970 s. The expansion of commercially important conifer plantations, which diminish habitat quality, and increasing agricultural depredation by Japanese Macaques, an apparent consequence of changes in human land use and rural demography, now influence management policies. More than 10,000 Japanese Macaques have been exterminated annually since 1998 as agricultural pests. Some of these monkeys are used for teaching purposes in medical and dental schools. Previously, the Japanese Macaque waslisted as Data Deficient on The IUCN Red List primarily because it is unclear if current population trends will assure its sustainability.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD5FFD1FA296FAFF736F6BF.taxon	materials_examined	Formosa (= Taiwan). Restricted by R. Kellogg in 1945 to “ Jusan (Ape’s Hill), Takao prefecture, Formosa ” (= Shou Shan, Kaohsiung, Hsien, Taiwan).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD5FFD1FA296FAFF736F6BF.taxon	discussion	M. cyclopis is a member of the fascicularis species group of macaques, including M. fascicularis, M. mulatta, and M. fuscata. It has the characteristic helmet-shaped and relatively long and narrow glans penis. M. cyclops, or its mainland progenitors, may represent an early stage in reduction oftail length in the fascicularis species group, and it presumably dispersed from the eastern Asian mainland opposite Taiwan to the island during drops in sea level correlated with glacial advances before 300,000 years ago. A molecular genetic study byJui-Hua Chu and colleagues suggested a divergence date for Taiwanese M. cyclopis and Japanese M. fuscata from mainland ancestral M. mulatta of 0 - 38 - 0 - 44 million years ago. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD5FFD1FA296FAFF736F6BF.taxon	distribution	Distribution. Endemic to Taiwan I, whereit is widely distributed in the E Mts.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD5FFD1FA296FAFF736F6BF.taxon	description	Descriptive notes. Head — body 47.5 - 65 cm (males, including captive specimens, 65 cm in one wild male) and 42 - 60 cm (females, including captive specimens, 44 cm in one wild female); tail 43.5 - 50 cm (males, including captive specimens) and 35 - 41 cm (females, including captive specimens); weight 6.3 - 18.5 kg (males, including captive specimens, 8 - 1 kg in one wild male) and 5.5 - 9.5 kg (females, including captive specimens). The mean head-body length of male Taiwanese Macaques exceeds that of females by 14 %. Dorsal pelage brightens from yellowish brown in the thoracic region to golden brown on the lumbosacral region. Crown is generally somewhat paler than thoracic region. Outer surfaces of limbs are about the same color as adjacent surfaces of the trunk, but they gradually become a little more grayish. Proximal one-third of dorsal surface of the tail is about the same color as the lumbosacral region, but the distal two-thirds is dark grayish brown to a clearly defined blackish; it is blacker in fully adult males than fully adult females. Undersurfaces of trunk, limbs, and tail are pale grayish. Skin of the thinly haired face is pinkish to reddish, while an indistinct fringe of somewhat elongated blackish hairs surrounds the face on the forehead and cheeks. Pelage of neonatesis dark gray to blackish and becomes about the same color as that of adults by ¢. 9 months of age.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD5FFD1FA296FAFF736F6BF.taxon	biology_ecology	Habitat. Broadleaf evergreen forest and, at upper elevations, mixed broadleaf needleleaf forest, and occasionally needle-leat and bamboo forest. The elevational range of the Taiwanese Macaque is 100 - 3400 m; about one-third of recorded localities are at elevations of 1000 - 1500 m, and only ¢. 5 % above 2500 m. The Taiwanese Macaqueis most frequently observed in the lower forest strata. It rests on exposed rocky surfaces on an uplifted coral reef formation in south-western Taiwan and enters cavesto sleep or apparently to keep cool. Its natural distribution appears not to extend to any small surrounding islands.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD5FFD1FA296FAFF736F6BF.taxon	food_feeding	Food and Feeding. The Taiwanese Macaque eats primarily fruits and seeds, although leaves and other plant parts from trees, shrubs, and herbs are included in the diet. Thirty-eight to 132 species of wild plants have been recorded in their diets at different sites. Crop raiding tends to occur at elevations below 1000 m, and at least 33 cultivated crops are known to be opportunistically exploited. Animal foods consumed by Taiwanese Macaques include crayfish, grasshoppers, cockroaches, termites, cicadas, beetles, caterpillars, flies, wasp nests, snails, tadpoles, lizards, and bird eggs.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD5FFD1FA296FAFF736F6BF.taxon	breeding	Breeding. Reproduction in the Taiwanese Macaque is strongly seasonal; mating frequency peaks in November — January and birth frequency peaks in March — June. A single young is born after an average gestation, in captivity, of 162 days. A twinning rate of 1 % was recorded at the south-western site of Shou Shan during five birth seasons. Females generally mature sexually at c. 3 - 5 — 4 - 5 years old, and they first give birth at 4 - 5 years old. Sexual maturation in males probably occurs by at least 5 - 6 years, at which time males emigrate from their natal groups. The mean length of the menstrual cycle in captive females has been recorded at 27 - 3 - 27 - 9 days and 29 - 4 days. Both singlemount and multimount ejaculations occur, and mating has been observed both on the ground and in the trees. In one captive group, copulation was preceded by rhythmic lip movements performed by the male and female facing each other. Conspicuous cyclical changes in female sexual skin are correlated with ovulatory cycles and may be restricted to the mating season. The hindquarters from the root of the tail to the back of the thighs become prominently red and swollen in subadult and young adult females; older females experience changes in redness but not tumescence. During the mating season, male facial and scrotal skin becomes intensely red and includes the appearance of a pair of narrow red streaks that extend laterally from the outer corners of the eyes. The known reproductive rate in natural populations is 0 - 45 - 0 - 95 births / sexually mature female / year; the mean annual reproductive rate is probably c. 0 - 7. Mean birth weight was 402 - 430 g in 111 captive births. Females in natural populations apparently nurse their infants for 6 - 12 months, and mean interbirth intervals of 13 - 4 months and 15 - 4 months were recorded at two study sites. Pelage coloration in neonatesis dark gray to blackish and becomes about the same as that of adults by ¢. 9 months old. Infanticide by immigrant males may be a cause of some infant mortality. A captive male in Japan reportedly lived for 39 years.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD5FFD1FA296FAFF736F6BF.taxon	activity	Activity patterns. Detailed information on activity patterns is available for only one population observed in Yushan National Park, central Taiwan, during 1987 - 1988: composite daily averages were 83: 2 % arboreal behavior and 16 - 8 % terrestrial behavior. Well-habituated groups of Taiwanese Macaques in Kenting, Shou Shan, and Fushan tend to move on the ground when traveling in disturbed areas and through trees when traveling in mature forests. Peak feeding timeis in the early morning, and a secondary peak occurs in the afternoon. The proportion of daylight hours spent foraging was 30 - 35 % and c. 60 % at two differentsites. Taiwanese Macaques sometimes sleep on the ground and in caves. They reportedly swim well and sometimes in the sea.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD5FFD1FA296FAFF736F6BF.taxon	biology_ecology	Movements, Home range and Social organization. Most groups are relatively small; a rough estimate of mean size in 121 groups surveyed was 12 - 2, with individual group numbers ranging from 2 - 10 to more than 20. Larger groups tend to be restricted to elevations below 1500 m. Troops are multimale-multifemale with resident lineages of female kin and males migrating from natal groups. A few transfers of adult females have also been observed. The pooled ratio of sexually mature males to sexually mature females varied from 1: 1 - 8 to 1: 6: 7 in eight natural populations, with a pooled sex ratio of 1: 2. All male groups including adults and juveniles have been observed at two localities and were loosely associated with mixed-sexed groups. Solitary monkeys, presumably males, have also been observed. The mean home range is ¢. 130 ha, and home ranges of neighboring groups may partly overlap. Local seasonal movements in response to temperature changes may occur in both lowland and upland populations.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD5FFD1FA296FAFF736F6BF.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Least Concern on The [UCN Red List. The Taiwanese Macaque is threatened mainly by habitat destruction. Local extirpation in the western lowlands due to intensive human settlement and agricultural exploitation had occurred by the mid- 19 " century, and logging continues to be a major threat. Taiwanese Macaques were hunted or trapped for food or medicinal purposes, including by aboriginal peoples, but hunting declined during the last two or three decades of the 20 " century, primarily as a result of conservation legislation. Taiwanese Macaques trapped as pests have been used for medical experimentation and teaching, and they were formerly trapped for export, mostly to Japan. An isolated population of Taiwanese Macaques, estimated at 1247 individuals in 2008, inhabits Shou Shan (Mt. Longevity) in Kaohsiung City, the largest city in southern Taiwan. Habitat degradation and provisioning have led to close contact and conflict between Taiwanese Macaques and people. Conflict between wildlife and tourists has gradually become a serious problem throughout Taiwan. Staff ofYushan National Park are training volunteers to educate tourists not to feed Taiwanese Macaques and how to interact with macaques on the new Central Cross-Island Highway. In 2001, the total extant population of Taiwanese Macaques was estimated to be ¢. 250,000 in ¢. 10,000 groups, but variability in these estimates was high.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD9FFDCFF2264A4F81DFC47.taxon	materials_examined	Kenya, Tana River, Mitole (2 ° 10 ’ S, 40 ° 10 ’ E).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD9FFDCFF2264A4F81DFC47.taxon	discussion	Some experts have included the taxa C. aghs and C. sanjei as subspecies of C. galeritus, while others give these taxa full species status. C. galeritus is broadly sympatric with Piliocolobus rufomitratus and Cercopithecus mitis albotorquatus; narrowly sympatric on the forest edges with Papio cynocephalus and Chlorocebus pygerythrus. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD9FFDCFF2264A4F81DFC47.taxon	distribution	Distribution. SE Kenya, endemic to riparian forest patches along the lower Tana River; an additional small population was recently discovered in the Tana River Delta.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD9FFDCFF2264A4F81DFC47.taxon	description	Descriptive notes. There are no specific measurements available. In his 1997 Field Guide to African Mammals, J. Kingdon provided the following estimates: head — body c. 50 - 63 cm (males) and c. 42 - 53 cm (females), tail ¢. 45 - 77 cm; weight ¢. 8 - 10 kg (males) and ¢. 5.6 - 5 kg (females). The adult male type specimen, as measured by W. Peters, had a head-body length of 49 cm and a tail length of 72 cm. Fur of the Tana River Mangabeyis long, loose, and wavy, being a somewhat inconspicuously speckled grayish-yellow or mushroom color above, with fluffy-haired, yellowish-white underparts. Limbs are not speckled, and forearms, hands, and feet are dark brown. Tail has a slight, pale tuft, particularly in adult males. Face and ears are black, cheeks and temples are whitish, and eyelids are bright white. There is a center parting on the crown, beginning immediately behind the forehead, with very long, darker hairs diverging from it on either side. These hairs become even longer back toward the middle of the crown. Male Tana River Mangabeys call with a “ whoop-rumble, ” which is typical of mangabey species. Adult females are smaller than males.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD9FFDCFF2264A4F81DFC47.taxon	biology_ecology	Habitat. Dry riparian gallery forests and adjacent patches of bush along the Tana River. The Tana River Mangabey is highly dependent on old growth forest. January and February are the hottest and driest months. Annual rainfall is 500 - 600 mm and is distributed bimodally: March — June (peaks in March-April) and October-December (peaks in November-December). The Tana River often changesits course, resulting in ox-bow lakes that have forest patches around them that are occupied by mangabey groups. Gallery forest depends on ground water. Changes in river course affect forest dynamics — patches along the old river bed dry up while new patches are established along new river courses. Sometimesthe forest is flooded for extended periods (canopy trees may die and fall), resulting in open patches in the canopy. As such, forests used by the Tana River Mangabey are characterized by a mosaic of habitats in different stages of succession.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD9FFDCFF2264A4F81DFC47.taxon	food_feeding	Food and Feeding. Tana River Mangabeys mainly eat fruits, especially figs, and hard nuts and seeds, with some leaves, flowers, and animal prey (including invertebrates and small vertebrates such as frogs and lizards). They spend ¢. 73 % of their feeding time on the ground or in the undergrowth (0 - 2 m high). At other times, they feed in trees up to and over 20 m high. During food scarcity (dry season), they spend more time foraging and travel farther.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD9FFDCFF2264A4F81DFC47.taxon	breeding	Breeding. Female Tana River Mangabeys most likely have a 30 day reproductive cycle. A single young is born after a gestation of 180 days. As in other mangabey species, females often exhibit post-conception swellings. Individuals may live up to 20 years. Birth season is in November — February. The interbirth interval is two years.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD9FFDCFF2264A4F81DFC47.taxon	activity	Activity patterns. Tana River Mangabeys are diurnal, terrestrial, and arboreal.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD9FFDCFF2264A4F81DFC47.taxon	biology_ecology	Movements, Home range and Social organization. The average size of a group home range of the Tana River Mangabey is ¢. 47 ha. The average daily movement is 1500 m (range 1000 - 2600 m). Tana River Mangabeys live in multimale-multifemale groups, averaging 27 individuals (range 6 - 62). Groups may have up to six adult males. Larger groups may be fusions of smaller units. The population density is less than 0 - 07 ind / km?.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD9FFDCFF2264A4F81DFC47.taxon	conservation	Status and Conservation. CITES Appendix I. Classified as Endangered on The IUCN Red List. The Tana River Mangabeyis listed as Class A in the African Convention on the Conservation of Nature and Natural Resources. It survives as a single population in an extremely small area, confined to a 60 km-long narrow strip of fragmented gallery forests. In 1976, the 175 km? Tana River National Primate Reserve was established to protect the Tana River Mangabey and the equally threatened Tana River Red Colobus (Piliocolobus rufomitratus). This reserve harbors ¢. 56 % of the extant groups of Tana River Mangabeys. Of the 44 % living outside of reserve, 10 % live in forests under management of the Tana Delta Irrigation Project, while the remainder is on Trust / Government land. Damming and irrigation cause changes to the natural flood regime of the Tana River and result in vegetation change and the loss of gallery forests. Forest clearing, habitat degradation, and fires are serious threats. The Tana River Mangabey (and the Tana River Red Colobus) depends on old growth forest, which is now found only in few and isolated fragments because of their constant destruction by humans (e. g. clearing for agriculture, dyke construction, burning, extraction of honey, building materials, and firewood). Manyfruiting trees important to Tana River Mangabeys are cut to make canoes and for building. A favored source of fruits — the palm, Phoenix reclinata (Arecaceae) — is cut to make palm wine. In 2008, the total population of Tana River Mangabeys was estimated at ¢. 1200 individuals. In 2001, 2070 individuals were counted in 59 groups. The Tana River Mangabey needs to be closely monitored given the insecurity of its remaining habitat and potential future threats.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD9FFDDFA2A6111FD1DFCD4.taxon	materials_examined	Congo-Brazzaville, Poste des Ouaddas, Oubangui-Congo confluence.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD9FFDDFA2A6111FD1DFCD4.taxon	discussion	There are fairly well separated dark and light morphs of C. agilis, with the former being much more common. Individuals tend to be larger in the western part ofits distribution. Albinism has been reported. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD9FFDDFA2A6111FD1DFCD4.taxon	distribution	Distribution. SE Cameroon, SW Central African Republic, N DR Congo (N of the Congo River, from Ubangi River to the Ituri Forest and the Semliki River), NE Gabon (E of the Ivindo River), and the N of the Republic of Congo (across the Sangha River). A single record from Equatorial Guinea was possibly a misidentification.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD9FFDDFA2A6111FD1DFCD4.taxon	description	Descriptive notes. Head-body 51 - 65 cm (males) and 44 - 55 cm (females), tail 56 — 76 cm (males) and 45 - 60 cm (females); weight 8.8 - 10.2 kg (males) and 4.3 - 6.2 kg (females). The Agile Mangabeyis a large, slender mangabey, with relatively long limbs, a long muzzle, and a tail that is longer than its body. Fur is short and fine, being brownish-olive to gray-olive (speckled with yellow) above, but the median dorsal zone tends to be darker. Underside (as far as the chin and including inner surfaces of limbs) is pale grayish-white and not speckled. Hands and feet of the Agile Mangabey are very dark brown, almost black. Tail becomes lighter distally, is light below, and sometimes ends in a whitish tuft. Face is black with pale (not white) eyelids, and cheeks are white. Crown is somewhat dark, nearly always with a whorl of hair often exposing a pale patch of skin. Males are much larger than females. Adult males produce a loud, long-distance call, a “ whoop ” followed by a rumble that can be heard from up to a kilometer away.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD9FFDDFA2A6111FD1DFCD4.taxon	biology_ecology	Habitat. Mostly seasonally flooded forest and rainforest near rivers. In contrast, Agile Mangabeys in Dzanga-Sangha Forest Reserve, Central African Republic, spend most of their time in terra firma-mixed forest and are never observed along the Mondika River or in swampy forest along the Ndoki River. As. in other capped mangabey species, the Agile Mangabey spends most ofits time on the ground and in the lower forest strata (0 — 10 m).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD9FFDDFA2A6111FD1DFCD4.taxon	food_feeding	Food and Feeding. Agile Mangabeys eat mainly fruits and seeds (up to 76 %), supplemented with underground plant parts, herbs, pith, shoots, and some animal prey (including young duikers, Cephalophus).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD9FFDDFA2A6111FD1DFCD4.taxon	breeding	Breeding. Female Agile Mangabeys have a menstrual cycle of ¢. 30 days. A single young is born after a gestation of c ¢. 6 months. Interbirth intervals average 21 months if the infant survives. As in other mangabey species, females develop conspicuous perineal swellings when approaching ovulation. Post-conception swellings are also likely.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD9FFDDFA2A6111FD1DFCD4.taxon	activity	Activity patterns. Agile Mangabeys are diurnal and semi-terrestrial.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD9FFDDFA2A6111FD1DFCD4.taxon	biology_ecology	Movements, Home range and Social organization. The home range size of a group of Agile Mangabeys at Dzangha-Sangha was ¢. 303 ha. Another group in Gabon used 198 ha. Daily movement at Dzanga-Sangha was 1155 m (range 390 - 1985 m). The group size was 8 - 22 individuals, but groups of 40 and more than 100 individuals have also been observed. Mean group size in the Gabon study was 10 - 5 individuals (7 - 18). Groups usually contain several adult males and females. Densities in Gabon were 6 - 7 — 12 - 5 ind / km?, but lower densities have been recorded elsewhere.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD9FFDDFA2A6111FD1DFCD4.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Least Concern on The [UCN Red List. The Agile Mangabeyislisted as Class B in the African Convention on the Conservation of Nature and Natural Resources. It is threatened by habitat loss caused by deforestation. It is also locally hunted for meat and persecuted for crop raiding. Studies are needed to assess the impact of hunting on the Agile Mangabey. It is known or presumed to occur in a number of protected areas: Minkebé National Park in Gabon; Dja Biosphere Reserve in Cameroon; Dzanga-Sangha Forest Reserve, Ngotto Classified Forest, and Dzanga-Ndoki and Mbaéré-Bodingueé national parks in the Central African Republic; Garamba National Park in DR Congo; and Nouabalé-Ndoki and Odzala national parks in the Republic of the Congo.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD8FFDDFFF4667EFB66F3A2.taxon	materials_examined	Democratic Republic of Congo, south of Congo River.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD8FFDDFFF4667EFB66F3A2.taxon	discussion	C. chrysogaster is ranked as a subspecies of C. galeritus by some experts. Monotypic. Distribution. Congo Basin in the DR Congo, S of the Congo River. Its precise N, S and E distributional limits are not fully known, but based on current records, the W limit is the Congo River, the N limit could be the Lulonga River, and the E limit is probably the Lomami River.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD8FFDDFFF4667EFB66F3A2.taxon	description	Descriptive notes. There are no specific measurements available. In his 1997 Field Guide to African Mammals, J. Kingdon provided the following estimates: head-body c. 52 - 66 cm (males) and c. 45 - 55 cm (females), tail ¢. 50 - 75 cm; weight c. 10 - 14 kg (males) and c. 6 - 8 kg (females). The Golden-bellied Mangabey is relatively robustly built, and its striking coloration and naked violet rump are similar to the Drill (Mandrillus leucophaeus). Upper body, crown, and outer limbs of the Golden-bellied Mangabey are a rich, dark, speckled reddish-brown or grizzled olive, which is in sharp contrast with creamy-yellow cheeks, throat, and inner surfaces of limbs. Chest and belly are bright golden orange, hence the common name. Cheek whiskers are long and swept back. Eyelids are pale but not white. Tail is speckled at the root only, tapered, and carried in a backward arc (unlike other mangabeys). Size differences between males and females are considerable.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD8FFDDFFF4667EFB66F3A2.taxon	biology_ecology	Habitat. Rainforest and swamp forest up to 500 m above sea level. Golden-bellied Mangabeys occur in seasonally inundated lowland rainforest and may occur in gallery forest; they are sometimes also recorded in secondary forest. They prefer the ground and understory. The Golden-bellied Mangabeyis still poorly known with regard to its behavior, ecology, and reproduction.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD8FFDDFFF4667EFB66F3A2.taxon	food_feeding	Food and Feeding. Golden-bellied Mangabeys mainly eatfruits and seeds, along with leaves, flowers, gums, and animal prey.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD8FFDDFFF4667EFB66F3A2.taxon	breeding	Breeding. There is no specific information available for this species in the wild, but in captivity female Golden-bellied Mangabeys reach menarche, begin cycling, and have their first perineal swelling at ¢. 2 - 5 years of age. First pregnancies occurafterfive years of age. A typical menstrual cycle lasts 33 days. The gestation period averages 165 - 180 days. Interbirth intervals are 17 - 23 months. As in other mangabey species, female Golden-bellied Mangabeys often develop a post-conception swelling. The Goldenbellied Mangabey can live 30 years in captivity.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD8FFDDFFF4667EFB66F3A2.taxon	activity	Activity patterns. Golden-bellied Mangabeys are diurnal and mainly terrestrial.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD8FFDDFFF4667EFB66F3A2.taxon	biology_ecology	Movements, Home range and Social organization. No systematic observations of Golden-bellied Mangabeys have been made. There are reports of group sizes of 15 - 35 individuals but also of groups with more than 100 individuals. They most likely live in multimale-multifemale groups.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD8FFDDFFF4667EFB66F3A2.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Data Deficient on The [IUCN Red List. The Golden-bellied Mangabey is listed as Class B in the African Convention on the Conservation of Nature and Natural Resources. It is intensively hunted and may be an agricultural pest in some areas. It may also be at risk from localized habitat loss The Golden-bellied Mangabey occurs in Salonga National Park, DR Congo. Further survey work is urgently needed to determine the limits of its distribution, its abundance, and current conservation status.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD8FFDDFFE569B8F6B1F41E.taxon	materials_examined	Tanzania, Sanje Waterfall, Mwanihana Forest.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD8FFDDFFE569B8F6B1F41E.taxon	discussion	Some experts have regarded C. sanjei as a subspecies of C. galeritus. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD8FFDDFFE569B8F6B1F41E.taxon	distribution	Distribution. S Tanzania, confined to Mwanihana Forest and the E slopes of the Udzungwa Mts. Descriptive notes. There are no specific measurements available. In his 1997 Field Guide to African Mammals, J. Kingdon provided the following estimates: head-body c. b 0 - 65 cm, tail ¢. 55 - 65 cm. Body masses of male and female Sanje River Mangabeys are c. 10 - 3 kg and c. 5 - 8 kg, respectively. Crown, back, limbs and tail are speckledgray. Hands and feet are darker, and tail has a whitish tuft; belly and chest are pale orange. Face is pale grayish, with pink around the eyes and on the nose, and eyelids are off-white. Crown hairs part along midline or form a slight whorl; hairs extending back and to the sides are longer, with shorter hairs protruding outward along the brow. Body skin is bluish-white. Skin on hands, feet, and earsis dark grayish. Paracallosal skin is bluish gray, with narrow pink callosities. Sanje River Mangabeys are clearly sexually dimorphic; adult males are larger than adult females.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD8FFDDFFE569B8F6B1F41E.taxon	biology_ecology	Habitat. Montane and submontane forest fragments, preferring riparian forest and scrub at elevations of 400 - 1600 m. The Sanje River Mangabey is often found in the lower understory and spends more than 50 % ofits time foraging on the forest floor. It frequently moves through and uses disturbed areas and mosaic habitat. Most of the information on ecology, behavior, and reproduction of the Sanje River Mangabey comes from a habituated group in the Mwanihana Forest studied by K. Mwamende.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD8FFDDFFE569B8F6B1F41E.taxon	food_feeding	Food and Feeding. The Sanje River Mangabey has a variable diet, feeding on plant material (mainly fruits and seeds), insects, millipedes, and small animal prey, including slugs, snails, chameleons, frogs, and freshwater crabs. About 50 % of the diet is made up offruit, 27 % seeds, 11 % insects and other invertebrates, 4 % leaves, 2 % flowers, and 2 % bark and gums. They also occasionally eat more than 20 species of fungi, buds, lichens, and roots (4 %).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD8FFDDFFE569B8F6B1F41E.taxon	breeding	Breeding. Sanje River Mangabeys mate throughoutthe year. Females develop prominent perineal swellings during receptive periods. Mean menstrual cycle length is 32 days, and the gestation period averages 173 days. Females give birth to a single young. As in other mangabey species, female Sanje River Mangabeys exhibit post-conception swellings, on average, 67 days after conception.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD8FFDDFFE569B8F6B1F41E.taxon	activity	Activity patterns. Sanje River Mangabeys are diurnal and semi-terrestrial. They sleep in clumps oftall trees. One study group regularly used eight different sites scattered through its home range.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD8FFDDFFE569B8F6B1F41E.taxon	biology_ecology	Movements, Home range and Social organization. Group sizes of Sanje River Mangabeys are c. 15 to more than 40 individuals, and they are multimale-multifemale. Males disperse, and femalesare philopatric. An habituated study group in Mwanihana Forest Reserve had at least 63 individuals, including ten adult males, 21 adult females, and three subadults. Mean daily movement ofthis group was 1760 m, and its home range was ¢. 240 ha. Home range and movement patterns differed significantly between seasons. During the dry season, the group moved larger distances between forest patches and traveled significantly faster than during the wet season. Groups of Sanje River Mangabeys sometimes split temporarily to travel and forage separately (fission-fusion). Estimates of 0 - 3 - 0 - 6 groups / km?, with a mean of 10 - 2 ind / group, have suggested 900 mangabeys in the 177 km? Mwanihana Forest. Males display loud vocalizations known as the “ whoop-gobble ” calls.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD8FFDDFFE569B8F6B1F41E.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Endangered on The [UCN Red List. The Sanje River Mangabeyis listed as Class B in the African Convention on the Conservation of Nature and Natural Resources. It is estimated that fewer than 1300 individuals remain in two main populations, some 85 km apart (one consists of five distinct subpopulations). Sanje River Mangabeys are occasionally caught in snares set by local hunters. Across its distribution, the Sanje River Mangabey is threatened by habitat loss and degradation because of agriculture, infrastructure development, fires, wood extraction, and hunting. They occur in three protected areas: Mwanihana Forest Reserve, Udzungwa Mountains National Park, and Udzungwa Scarp Forest Reserve in Tanzania. Efforts to expand the boundaries of Udzungwa Mountains National Park to improve the level of protection in the current Udzwunga Scarp Forest Reserve have been unsuccessful. Demographic surveys are needed. There is a need to enforce laws prohibiting hunting in forest reserves and increase prevention of habitat alteration.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD8FFDEFAEB6937FB6FF26D.taxon	materials_examined	Afrique occidentale.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD8FFDEFAEB6937FB6FF26D.taxon	discussion	In his 1958 review of the zoogeography of West Africa, A. Booth reported the possibility of distributional overlap and hybridization of C. atys and C. lunulatus in a small area between the Sassandra and Nzo rivers. Complete albinism has been reported. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD8FFDEFAEB6937FB6FF26D.taxon	distribution	Distribution. SW Senegal, Guinea-Bissau, W & SE Guinea, Sierra Leone, Liberia, and W Ivory Coast (E boundary is the Nzo-Sassandra river system).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD8FFDEFAEB6937FB6FF26D.taxon	description	Descriptive notes. Head-body 43 - 6 - 58: 0 cm (males) and 47.4 - 53.0 cm (females), tail 52 - 60 cm (males) and 53.0 - 64.5 cm (females); weight 9.5 - 11.4 kg (males) and 5.6 - 7.0 kg (females). The grayish appearance of the Sooty Mangabey gives it its common name. It is generally smoky iron-gray above and paler ventrally, with darker hands and feet. Some individuals show a trace of a darker dorsalstripe. Face is dark grayish-pink, with a black muzzle and ears, and cheek whiskers are elongated and lighter than the body. There is generally no whorl on the crown. Nuchal markings are weakly expressed. The Sooty Mangabey is somewhat less sexually dimorphic than other mangabeys.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD8FFDEFAEB6937FB6FF26D.taxon	biology_ecology	Habitat. A variety of primary and secondary forest types including dry, swamp, mangrove, and gallery forest in the Guinean forest zone. The Sooty Mangabey is often found in estuaries among the root systems of mangroves. It also occurs in woodland savanna in Guinea and tropical moist evergreen rainforest in Tai Forest National Park, Ivory Coast. Groups occasionally raid farms and rice paddies. In the absence of hunting, Sooty Mangabeys are tolerant of some degree of habitat degradation.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD8FFDEFAEB6937FB6FF26D.taxon	food_feeding	Food and Feeding. Sooty Mangabeys eat mainly fruits, seeds, and nuts, along with some animal prey. In the Tai Forest, diets contain 68 % fruits and seeds and 26 % invertebrates. Elsewhere, plants make up a higher proportion of their diets (e. g. 98 - 7 % plant foods and only 1 -: 3 % animal foods). Of the 98 - 7 % plant foods, the vast majority was fruit, with small amounts of leaves, flowers, and miscellaneous plant parts. The Sooty Mangabey regularly eats frog eggs.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD8FFDEFAEB6937FB6FF26D.taxon	breeding	Breeding. In captivity, female Sooty Mangabeys have their first perineal swelling at 2: 5 - 3 - 3 years of age, and their first births usually occur at c. 4 - 7 years of age, which is most likely earlier than in the wild. Captive females have a mean 34 - 5 day ovarian cycle, with receptivity signaled by a conspicuous, pink genital swelling. The interbirth interval in captivity averages 1 - 4 years. Post-conception swellings have also been reported, which peak 49 days after conception. Usually a single offspring is born after a gestation of 167 days. Instances of infanticide have been reported. In the wild, Sooty Mangabeys have discrete and distinct mating seasons. Females utter copulation vocalizations during intercourse. Usually the alpha male of the group tries to monopolize copulations, but sneak copulations are common. During common group encounters in the wild, receptive females have been observed copulating with non-resident males and solitary non-group males. Individuals may live up to 18 years.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD8FFDEFAEB6937FB6FF26D.taxon	activity	Activity patterns. Sooty Mangabeys are diurnal and mainly terrestrial. It moves and forages mainly on the ground but will also use the forest canopy. In the Tai forest, 38 - 8 % of the Sooty Mangabey’s time is spent feeding, 24 - 5 % foraging 18: 5 % resting, 10 - 3 % traveling, and 7 - 9 % engaging in social activities.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD8FFDEFAEB6937FB6FF26D.taxon	biology_ecology	Movements, Home range and Social organization. The home range of the Sooty Mangabeyis estimated at 4 - 6 - 5 km *. Home ranges are known to overlap extensively in the wild. Intergroup encounters are common and typified by avoidance, each group ignoring the other or engaging in aggressive interactions. Daily movements average 2: 2 km. Sooty Mangabeys live in large groups of up to 120 individuals, with multiple males and multiple females. Two categories of males occur in groups in the Tai forest: some males are full-time residents, while others are more transient, alternating several weeks of residence in the group and several weeks of absence. Male Sooty Mangabeys disperse from their natal group, while females tend to stay, forming matrilines of closely related females. Males and females each establish linear dominance hierarchies, with adult males usually outranking females.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFD8FFDEFAEB6937FB6FF26D.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Vulnerable on The IUCN Red List. The Sooty Mangabey is listed as Class B in the African Convention on the Conservation of Nature and Natural Resources. Habitat destruction and hunting are the two most profound threats to the survival of the Sooty Mangabeys. It is known to occur in several protected areas: Tai Forest National Park and Monogaga Classified Forest in Ivory Coast; Sapo National Park and North Loma, Gola, and Grebo national forests in Liberia; Outamba-Kilimi National Park and Tiwai Island Wildlife Sanctuary in Sierra Leone; and Diécké Forest Reserve and Massif du Ziama Biosphere Reserve in Guinea. The Sooty Mangabey does not appear to be common in Senegal and Guinea-Bissau. In the absence of hunting, the Sooty Mangabey used to be relatively widespread in farmbush and secondary forest in Sierra Leone.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDBFFDEFA2D640CF802F554.taxon	materials_examined	Ghana, River Boutry.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDBFFDEFA2D640CF802F554.taxon	discussion	C. lunulatus is ranked as a subspecies of C. atys by some experts. This species was also formerly regarded as a subspecies of C. torquatus. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDBFFDEFA2D640CF802F554.taxon	distribution	Distribution. E & NE Ivory Coast (from the Nzo-Sassandra river system to the Volta River), SW Burkina Faso, and SW Ghana. It is believed to have been extirpated in the region between the Sassandra and Bandama rivers, Ivory Coast.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDBFFDEFA2D640CF802F554.taxon	description	Descriptive notes. Head — body 52 - 73 cm (males) and 47.4 - 51 cm (females), tail 67 - 5 — 74 cm (males) and 68 cm (one female); weight 7.1 - 11.8 kg (males) and 3.9 - 6.3 kg (females); all measurements from captive individuals. Although similar in their habits, male White-naped Mangabeys are larger than Sooty Mangabeys (C. atys). Female White-naped Mangabeys and Sooty Mangabeys are similar in size. Back, tail, outer limbs, hands, and feet of White-naped Mangabeys are a brownish smoky-gray, with a well-expressed dorsal stripe. Underside, chest, and throat are white. Face is grayish pink but darker than that of Sooty Mangabeys. There is a whorl on the crown, and a white half-moon shaped patch (bordered with black) across the nape, which resulted in its scientific name. Males are considerable larger than females.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDBFFDEFA2D640CF802F554.taxon	biology_ecology	Habitat. A variety of primary and secondary forest types, including dry, swamp, mangrove, and gallery forest. It rarely ascends beyond the lowest level of the canopy. Little is known about the White-naped Mangabey, but it most likely shares many features of its behavior and ecology with its sister species, the Sooty Mangabey.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDBFFDEFA2D640CF802F554.taxon	food_feeding	Food and Feeding. White-naped Mangabeys eat mainly fruits, along with seeds, other plant material, and some animal prey.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDBFFDEFA2D640CF802F554.taxon	breeding	Breeding. Female White-naped Mangabeys probably have a 30 day reproductive cycle, and they develop a conspicuous perineal swelling around ovulation. A single young is born after a gestation of ¢. 170 days.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDBFFDEFA2D640CF802F554.taxon	activity	Activity patterns. White-naped Mangabeys are diurnal and mainly terrestrial.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDBFFDEFA2D640CF802F554.taxon	biology_ecology	Movements, Home range and Social organization. The home range size of the Whitenaped Mangabey is c. 200 ha. Population densities can reach 5 ind / km? Groups of 3 - 58 individuals have been observed, most likely comprising several adult males and females.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDBFFDEFA2D640CF802F554.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Endangered on The [UCN Red List (as C. atys lunulatus). The White-naped Mangabey is listed as Class B in the African Convention on the Conservation of Nature and Natural Resources. It is relentlessly hunted for the bushmeat trade. The White-naped Mangabey has a restricted, patchy distribution and is not known to be abundant anywhere. It may still occur in seven protected areas: Ankasa Game Reserve and Dadieso and Yoyo River forest reserves in Ghana and Comoé and Marahoué national parks and Dassioko and Niegre forest reserves in Ivory Coast. It is also found in the Warigué protected area in Burkina Faso. The White-naped Mangabey is the mangabey in Bia and Nini-Suhien national parks and Krokosua Hills Forest Reserve in Ghana. Surveys in Ivory Coast confirmed its presence only in Tanoé Community Forest in the extreme south-eastern part of the country and Como ¢ National Park, where it is threatened by civil conflict and hunting. The TUCN / SSC Primate Specialist Group listed the White-naped Mangabey as one of the “ World’s 25 Most Endangered Primates ” in 2000 - 2006 because of its almost complete elimination from eastern Ivory Coast and western Ghana.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDBFFDFFA366FE2FC47F551.taxon	materials_examined	Ethiopia (= West Africa).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDBFFDFFA366FE2FC47F551.taxon	discussion	This species is monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDBFFDFFA366FE2FC47F551.taxon	distribution	Distribution. Patchily in forests not more than 350 km inland from the coast in S Nigeria (W limit in forests near Dahomey Gap), W Cameroon, W Equatorial Guinea, W Gabon, and SW of the Republic of Congo (as far S as Conkouati-Douli National Park); there have been unconfirmed reports of its occurrence in Benin, butif it ever did occur there, it may now be extirpated, as was a former population near the Benue River in Nigeria.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDBFFDFFA366FE2FC47F551.taxon	description	Descriptive notes. Head-body 46 - 67 cm (males) and 42 - 59 cm (females), tail 50 - 76 cm (males) and 46 - 64 cm (females); weight 8 - 125 kg (males) and 5 - 8 kg (females). The Red-capped Mangabey is long-tailed and long-legged. It is dark slate-gray above, with a sharply demarcated white underside (including inner surfaces of limbs, chin, neck, and cheeks). Face is dark with striking white eyelids. There is a conspicuous dark red patch on the crown, outlined by a white collar and temporal line. Tail has a white tuft and is frequently held so that the white tip comes forward over the head. Males are much larger than females.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDBFFDFFA366FE2FC47F551.taxon	biology_ecology	Habitat. Primary and secondary swamp and mangrove forest, and rainforest and gallery forest. The Red-capped Mangabey can also be found in young secondary forests and around cultivated areas.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDBFFDFFA366FE2FC47F551.taxon	food_feeding	Food and Feeding. Red-capped Mangabeys eat mainly fruits, seeds, and palm nuts, along with leaves, flowers, bark, gums, eggs, and animal prey (including invertebrates, reptiles, and small mammals). Fruits and seeds from canopy trees make up more than 80 % of their diet in the Campo Reserve in Cameroon. Often hard objects constitute a substantial part of the diet, such as seeds of the bitter bark tree or Liberian cherry (Sacoglottis gabonensis, Humiriaceae).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDBFFDFFA366FE2FC47F551.taxon	breeding	Breeding. In captivity, female Red-capped Mangabeys have a 26 - 35 day reproductive cycle, with receptivity signaled by a conspicuous pink genital swelling. As in other mangabey species, females regularly develop post-conception swellings. A single offspring is born after a gestation of 168 days (range 160 - 182 days). Neonates are almost hairless, and the red cap is not evident. Juveniles are much darker than adults. Individuals may live up to 27 years.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDBFFDFFA366FE2FC47F551.taxon	activity	Activity patterns. Red-capped Mangabeys are diurnal, arboreal, and terrestrial.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDBFFDFFA366FE2FC47F551.taxon	biology_ecology	Movements, Home range and Social organization. The Red-capped Mangabey seems to travel predominantly on the ground, but it mainly uses the middle and upper canopies when feeding (Sette Cama, Gabon, 44 % on the ground). In the Campo Reserve, a group used 250 ha during 15 months. Home ranges overlap with other groups. Daily movement is ¢. 1000 m. Group size is 20 - 60 individuals, including several adult males. In Sette Cama, a group of 60 individuals frequently separated into subgroups (fission) during foraging. Group members keep in contact by calls. Red-capped Mangabeys are often found in polyspecific association with species of Cercopithecus and, interestingly, Mandrills (Mandrillus sphinx), even though they evidently occupy a similar ecological niche. The social organization of Red-capped Mangabeys is probably similar to other species of capped mangabeys (Cercocebus); they live in multimale-multifemale groups with linear dominance hierarchies among males and females. Normally females remain in their natal groups, and males disperse to other groups.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDBFFDFFA366FE2FC47F551.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Vulnerable on The IUCN Red List. The Red-capped Mangabeyislisted as Class B in the African Convention on the Conservation of Nature and Natural Resources. It is threatened by habitat loss and hunting for meat throughout most ofits distribution. It is regarded as an agricultural pest in some areas. Although Red-capped Mangabeys seem locally abundant, they now appear to be absent in areas with even low to medium hunting pressure. Red-capped Mangabeys occurs in 18 protected areas: Banyang-Mbo Wildlife Sanctuary, Douala-Edéa Wildlife Reserve, and the Campo-Ma’an and Korup national parks in Cameroon; Monte Alen National Park in Equatorial Guinea; Loango, Mayumba, Moukalaba-Doudau, and Pongara national parks and Sette-Cama Reserve in Gabon; Okomu and Cross River national parks and Igangan Ohosu, Oluwa, and Omo forest reserves in Nigeria; and Conkouati-Douli National Park in the Republic of the Congo.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDAFFD8FFEB6EAAFA9EFD83.taxon	materials_examined	Borneo. Restricted by E. Delson and P. Napier in 1976 to “ Cameroon, Bitye, Dja River. ”	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDAFFD8FFEB6EAAFA9EFD83.taxon	discussion	Molecular studies suggest that there are perhaps two distinct populations of M. sphinx. A study by P. T. Telfer and colleagues in 2003 indicated that the Ogooué River in Gabon bisects their distribution, separating them into two distinct populations: Cameroon and northern Gabon, and southern Gabon. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDAFFD8FFEB6EAAFA9EFD83.taxon	distribution	Distribution. W Central Africa, from S Cameroon (S of the Sanaga River) to mainland Equatorial Guinea, W Gabon (Ivindo and Ogooué rivers limit its distribution to the E), and SW Republic of the Congo (S to the Kouilou River and down to the Congo River). It is not known to occur east of the Dja River in Cameroon, and it does not occur in the forests of SE Cameroon.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDAFFD8FFEB6EAAFA9EFD83.taxon	description	Descriptive notes. Head-body 62 - 110 cm (males) and 55 - 67 cm (females), tail 7 - 10 cm (males) and 5 - 9 cm (females); weight 18 - 33 kg (males) 11 - 13 kg (females). Mandrills and Drills (M. leucophaeus) are the largest living monkeys and certainly among the most physically impressive. Fur is long and thick and grayish-brown dorsally, with a sulphuryellow underside. Tail is short and invariably held erect. Sexes differ remarkably in size; female body mass is 36: 5 % of male body mass. Males have a proportionately bigger head than females, a heavy mane, massive muzzle, and formidable upper canines. Besides their enormous build, Mandrills have remarkable coloration, best exemplified in the deeply ridged, brilliantly hued snout and equally luminous rump of the dominant male. The bridge of the nose and lips are a rich, lacquered scarlet, while sides of the nose are pale blue with traces of purple in the grooves. There are fringes of whitish orange fur on cheeks and white behind the ears and a small crest atop the head. When a male has attained alpha status, he undergoes physical changes and becomes more brightly colored and has a higher testosterone level. With a loss ofstatus, the reverse changes occur. Both sexes sport a yellowish-orange beard. The rump also distinguishes mature males; it is adorned with a broad patch of blue, red, and violet skin, set against prominent reddish genitalia. Mandrills, like Drills, have a sternal gland. Both sexes use this gland to scent-mark by rubbing their chests on a substrate or object. The functions of scent marking are not clear. It may help reinforce dominance status, it may play a role in mate choice because odors can encode information on sex, rank, and genotype, or it may be used to orient within the home range.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDAFFD8FFEB6EAAFA9EFD83.taxon	biology_ecology	Habitat. Moist evergreen rainforest stretching 100 - 300 km inland from the Atlantic coast, semi-montane forest, and dense secondary forest. Mandrills are known to occur in forest fragments in savanna and use plantations, particularly during the dry season. There are two rainy and two dry seasons of roughly equal durations in the distribution of the Mandrill; the timing of these seasons differs north and south of the Equator.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDAFFD8FFEB6EAAFA9EFD83.taxon	food_feeding	Food and Feeding. Mandrills are omnivorous, and their diets are diverse, including fruits, buds, leaves, roots, fungus, seeds, and animal prey (insects, crabs, fish, amphibians, lizards, birds, and small mammals up to the size of duikers). They prefer fruits when they are available. In their primary forest habitat, fruiting of trees and lianas is irregular, leading to periodic fruit shortages. When this occurs, Mandrills are highly reliant on having abundant herbaceous plants to eat. When food is scarce (during and at the end of the dry season), they also raid crops. They forage mainly less than 5 m off the ground. Males spend most of their time on the ground, but females and young will forage in trees.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDAFFD8FFEB6EAAFA9EFD83.taxon	breeding	Breeding. There is no specific information available for this species in the wild, but in captivity or semi-captivity, female Mandrills have, on average, a 33 day menstrual cycle. Onset of the receptive period is signaled by genital swelling and reddening that reaches its peak around ovulation. The privilege of mating appears to be mostly restricted to alpha males. There is usually no set breeding season, but births are most frequent in December — April. Usually a single young is born after a mean interbirth interval of less than two years. The gestation period lasts 179 - 182 days. Infants have a pinkish face, silvery gray pelage, white hair on limbs, and a black cap. Juvenile pelage is similar to adults, and they have a dull blue snout and a buffy beard — characteristics that most females retain for life. Adult female Mandrills can be quite pink in the face, but not as much as the males. Females are sexually mature at 4 - 5 years old, and males at 5 - 7 years old. Individuals have been known to live for more than 45 years in captivity.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDAFFD8FFEB6EAAFA9EFD83.taxon	activity	Activity patterns. Mandrills are diurnal and mainly terrestrial.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDAFFD8FFEB6EAAFA9EFD83.taxon	biology_ecology	Movements, Home range and Social organization. The home ranges of Mandrills can be 30 - 50 km? and groups travel up to 15 km / day when foraging. The social organization of Mandrills has historically been unclear, with contradictory observations because they live in dense rainforests and are difficult to follow and observe. Most data have been derived from a semi-free-ranging colony of Mandrills in Gabon, where stable social groups consist of several adult males and females and their offspring. Early reports of wild Mandrills, based on only a few observations, suggested a multilevel social structure similar to Hamadryas Baboons (Papio hamadryas) and Geladas (Theropithecus gelada), with units of one male and several females as the basal social grouping. Nevertheless, more recent field studies have provided a different picture. Mandrills aggregate into extremely large groups of up to several hundred individuals, termed “ hordes. ” These large groups seem to remain cohesive and stable (e. g. Lopé National Park, Gabon), but they may split up into smaller groups temporarily. Most unusual for other species in the tribe Papionini, adult male Mandrills at Lopé reside in these large groups only during the mating season and are otherwise solitary.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDAFFD8FFEB6EAAFA9EFD83.taxon	conservation	Status and Conservation. CITES Appendix I. Classified as Vulnerable on The IUCN Red List. The Mandrill is listed as Class B in the African Convention on the Conservation of Nature and Natural Resources. It is threatened by habitat loss and hunting, and it is sometimes regarded as a pest and killed. Total population size is unknown, but it has undoubtedly declined in recent years. Mandrills occur in 14 protected areas: Campo Ma’an National Park in Cameroon; Monte Alen National Park in Equatorial Guinea; Minkebé, Crystal Mountain, Lopé, Moukalaba-Doudau, Waka, Mayumba, Ivindo, and Birougou national parks, Wonga-Wongue Presidential Reserve, L. ékédi Sanctuary, and Mont Iboundji Sanctuary in Gabon; and Conkouati-Douli National Park in the Republic of the Congo. Several of these need immediate protection, both legal and practical, against logging and hunting.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDDFFD8FF2367DFF973F3F5.taxon	materials_examined	Africa.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDDFFD8FF2367DFF973F3F5.taxon	discussion	Bioko Island is just 30 km offshore and believed to have been connected to the mainland as recently as 10,000 years ago. Two subspecies recognized.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDDFFD8FF2367DFF973F3F5.taxon	distribution	Subspecies and Distribution. M. l. leucophaeusF. Cuvier, 1807 — fromtheCrossRiverinSENigeria, EtotheSanagaRiveranditstributarytheMbamRiverinWCameroon; inlandfromthecoast, the distributionislimitedtotheNbythederivedsavannaboundary. M. l. poensis Zukowsky, 1922 — Bioko I.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDDFFD8FF2367DFF973F3F5.taxon	description	Descriptive notes. Head-body 72 - 83 cm (males) and 45.5 - 50 cm (females), tail 8 - 6 — 12: 5 cm (males) and 5.9 - 4 cm (females); weight 14.5 - 27 kg (males) and 6.5 - 12 kg (females). The Drill has extreme sexual dimorphism; weight of adult males is more than twice that of adult females. The distinctive smooth, bare, black face, with paranasal ridges, in both sexes is moon-shaped and greatly enlarged in males. Edges of the black face are sharply demarcated by white skin, giving a dramatic, mask-like appearance. Fully adult males bear a bright red strip between the black lower lip and white chin. Drills have white chin hairs without beard in adults and whiskers in males. Ears are naked, and black; bare skin behind ears is white and prominently visible in males. Forehead and crown are dark brown to blackish. Males have a semi-erectile crest, and the mantle of the neck and shoulder hair look like a short mane. Body hairis light brown to tawny gray, ventral hairs paler to white, with a faint brown dorsal stripe in adult males. Lower limbs are darker. Both sexes have a sternal scent gland; adult males have a pectoral tuft that holds aromatic secretions. Hands and feet are black. Ventral skin is white, and dorsal skin is blue-gray. Adult male Drills have a spectacular rump coloration exposed by sparse hair, displaying a deep spectrum from hot pink to lavender and blue, depending on age and condition. In both sexes, ischial callosities are pale pink. Skin around the anus and female sexual skin to the tail are red to pink. Adult males have a prominent heavy jaw and impressive canines; upper canines may exceed 5 cm. Females and males up to four years of age appear similar. Neonates have white to pink skin on face, ears, hands, and feet; they have silvery-gray hair and black “ infant caps. ” Infant pelage is replaced gradually over 3 - 6 months to eventually resemble color of adult females. Face darkens to black by 4 - 6 months. The “ Bioko Drill ” (M. I. poensis) is smaller and less sexually dimorphic than the Mainland Drill (M. I. leucophaeus). Relative to the Mainland Drill, the skull of the Bioko Drill is shorter and broader, overall stature is slightly less, and hair color may be less tawny and more blackish.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDDFFD8FF2367DFF973F3F5.taxon	biology_ecology	Habitat. Closed forest from coastal lowland to transitional and semi-montane forest further inland. These areas give way to forest-savanna mosaic and high-elevation grassland with gallery forest where Drills are absent. In the 1930 s, I. T. Sanderson collected Drills in mixed grassland — gallery forest of Cameroon’s Bamenda Highlands, but these forests have now been largely eliminated. Across their limited distribution, there is a single long dry season in December — March; Bioko Island is wetter.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDDFFD8FF2367DFF973F3F5.taxon	food_feeding	Food and Feeding. Drills are omnivorous, opportunistic, wide-ranging foragers. Fruits and seeds dominate their diets; other plant material includes blossoms, leaves, stems and pith, forbs, grass blades, corms, roots, tubers, and bark. Animal matter includes adult and larval insects and other invertebrates such as crabs and snails, bird and reptile eggs, small fish, reptiles, and amphibians. Drills will kill small birds and mammals; feathers, hair, and bone have been found in feces in the wild. They also eat mushrooms. Drills forage mostly on the ground, with some individuals feeding in the trees above. Younger animals are more aptto forage in the trees because their greater mobility and smallersize allow them to gain access to small branches, thereby reducing competition. Drills are avid diggers, scraping the forest floor, shifting leaflitter and other detritus, and breaking open rotten logs, branches, and insect mounds Their foraging leaves a distinctive sign of a group’s passing. Drills readily enter shallow water to capture tadpoles and other aquatic foods, and they dig up marine turtle eggs on Bioko Island beaches. Studies in Korup National Park, Cameroon, found that fruits and seeds comprised more than 82 % of the diet by weight, depending of the season. Insects or other animal matter was 5: 5 %.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDDFFD8FF2367DFF973F3F5.taxon	breeding	Breeding. Female Drills exhibit a prominent sexual swelling and cycle throughout the year; mean cycle length is 33 days. A single infant is born; twins are very rare. The mean gestation period is 176 days; 72 % of births occur in the rainy season with nearly 50 % of births in the two peak rainy months (June and July). A female may reproduce every year and will not actively wean her infant until her next infant is delivered. Females enter puberty at three years of age and typically beartheir first offspring at four years. Males are sexually mature at five years of age and will double in size before reaching full maturity at 8 - 10 years. These reproductive data are derived from more than 400 births at the Drill Ranch project in Nigeria, where Drill groups live in multi-hectare, electric fenced enclosures of natural habitat. The dominant male is believed to sire most offspring in his group.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDDFFD8FF2367DFF973F3F5.taxon	activity	Activity patterns. By day, a Drill group moves through the forest, foraging on the ground and in the lower levels of trees. They may rest or otherwise reduce travel and active foraging in the early afternoon. Before dusk, the group ascends a single large tree or multiple trees in close proximity, where some feeding and movements continue. There is no distinctive male loud call at this time, but contact calls between mothers and their young are heard. By nightfall, each Drill has selected a sleeping site from 10 m upward to tree crowns and the group is quiet. Infants sleep with their mothers. By dawn, contact calls and movements commence followed by urination and defecation; soon after, the group descends and assembles to begin its day’s activities.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDDFFD8FF2367DFF973F3F5.taxon	biology_ecology	Movements, Home range and Social organization. It is unclear if groups of Drill maintain a fixed home range or move across a larger area in semi-nomadic, seasonal patterns. Much anecdotal evidence and some field observations indicate Drills roam the forest in semi-regular, annual patterns, perhaps following fruiting trees or other food and water availability. Study sites in Korup and Ebo national parks have found Drills throughout the year. Drills may range more widely in less resource rich areas, such as higher elevation or marginal forests. Social organization data are limited and somewhat speculative. Drills live in multimale-multifemale groups with a dominant male. Typical group sizes are 15 - 30 individuals, with roughly a 1: 2 ratio offully adult males to the rest of group. “ Supergroups ” of 100 individuals or more may be temporary fusions of typical, dominant-male units. Supergroup formation may be seasonal and function as an opportunity for individuals to transfer between groups. Solitary Drills are uncommon, with only adult males observed alone.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDDFFD8FF2367DFF973F3F5.taxon	conservation	Status and Conservation. CITES Appendix I. Classified as Endangered on The IUCN Red List, including both subspecies. The Drill is listed as Class B in the African Convention on the Conservation of Nature and Natural Resources. With a small natural distribution of less than 50,000 km? in parts of three countries, the Drill’s statusis precarious. Within this distribution, less than 20,000 km? are usable for Drills and fragmented into at least eleven “ habitat islands ” on the mainland and two on Bioko Island. These restricted areas have a relatively high human density and several large urban centers that fuel the demand for bushmeat. Hunters may use dogs, whereby a group of Drills can be treed and several of them shot in a single hunting encounter; verified reports of up to 25 such kills exist. Commercial hunting for the last 30 years has greatly reduced the numbers of Drills. Habitat is lost primarily to farming but also to illegal logging in Nigeria and to farming and industrial logging in Cameroon. Large areas were cleared for palm, banana, and rubber plantations in Cameroon in early colonial days, and a recent upsurge in palm plantation proposals threaten important core habitat of Drills and other wildlife. Roads and further fragment habitat and ease access for hunters, bushmeat traders, farmers, and timber poachers. Drills are well protected by law from hunting and trade in all three countries where they occur, but enforcementis poor or absent. In Nigeria, most remaining habitats are technically protected in the two discontinuous divisions of the 3440 km * Cross River National Park; additional protected habitats include the smaller Afi Mountain Wildlife Sanctuary (100 km? *) and Mbe Mountain community forest (80 km?). Cameroon contains ¢. 75 % of the Drill’s remaining habitat. Korup National Park covering 1259 km? is the earliest protected area and is the core of the Drill’s current distribution. New and proposed protected areas include Takamanda, Ebo, Mount Cameroon, and Bakossi national parks. Banyang-Mbo Wildlife Sanctuary and numerous forest reserves nominally protect additional habitat but do not control hunting. Most protected areas form landscape blocks with contiguous forest reserves and community forests, and some across the Nigeria — Cameroon border. On Bioko Island, Drill habitat is protected in Basilé National Park (330 km?) and Luba Caldera Scientific Reserve (510 km?); these areas cover more than 40 % ofthe island. Drills are protected in part of Luba Caldera but may already be extirpated in Pico Basilé. Despite existing and planned habitat protection in all countries where the Drill occurs, the control of hunting is inadequate. As groups and subpopulations shrink and become increasingly isolated from one another, Drills may be unable to form supergroups and may lose viability. Population estimates are less than 10,000 individuals and possibly as low as 4000. A large managed population of captive Drills living in semi-wild conditions in Nigeria is prepared for group reintroduction to reinforce local subpopulations if needed.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDDFFD9FA306804F60BFC1E.taxon	materials_examined	Democratic Republic of Congo, Mayombe.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDDFFD9FA306804F60BFC1E.taxon	discussion	The genus Lophocebus was separated from Cercocebus and subsequently split into L. albigena and L. aterrimus. In 2007, C. P. Groves elevated three subspecies of L. albigena to full species, based on morphological characters and color of the shoulder mantle: gray in L. albigena, reddish-brown in L. osmani, and brown to straw-colored in L. johnstoni. Groves also recognized a fourth species (L. ugandae) based on skull morphology and smaller size. Some experts consider the grayer, coastal Cameroon population to be a distinct subspecies (L. a. zenkeri), although differences would seem to be more likely clinal in nature. Distributions of L. albigena and L. osmani might overlap in south-eastern Cameroon, and L. albigena possibly hybridizes with L. osmani in the Dja Biosphere Reserve, Cameroon, but is probably geographically separated from L. johnstoni by the Ubangi River and from L. aterrimus by the Congo River. Albinism is known in this species. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDDFFD9FA306804F60BFC1E.taxon	distribution	Distribution. S Cameroon (S of Sanaga River), SW Central African Republic (W of Ubangi River), Equatorial Guinea, Gabon, Republic of the Congo (E to Ubangi River).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDDFFD9FA306804F60BFC1E.taxon	description	Descriptive notes. Head-body 45 - 73 cm (males) and 50 - 56 cm (females), tail 73 — 100 cm (males) and 67 - 80 cm (females); weight 6.1 - 9 kg (males) and 4.7 - 6.5 kg (females). As in other species of crested mangabeys (Lophocebus), fur of the Gray-cheeked Mangabey is soft and long, being generally blackish-brown. A long, pale mantle on shoulders (more prominent in males) runs continuous with rather thin, grayish cheek whiskers. There is a short, partially upright tuft on the crown, and eyebrows tend to stand upright in two little “ horns. ” Tail is long and ragged, and it is normally carried straight up with the tip arched forward over the back. Face is black. Female Graycheeked Mangabeys have more slender builds than males and develop a conspicuous pink swelling when sexually receptive.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDDFFD9FA306804F60BFC1E.taxon	biology_ecology	Habitat. Low-elevation primary and secondary rainforest, gallery forest, and some swamp forest. The Gray-cheeked Mangabey is mosttypically found in dense evergreen forest, where it prefers the middle and upperlayers of the forest canopy.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDDFFD9FA306804F60BFC1E.taxon	food_feeding	Food and Feeding. As in otherspecies of crested mangabeys, the diets of Gray-cheeked Mangabeys consist mainly offruits (30 - 60 %) and seeds (29 - 36 %). Theyalso eat young leaves, bark, flowers, and animal prey (invertebrates and small vertebrates). Animal prey makes up 27 % of the diet at Dja, Cameroon, 13 % at Mondika, Central African Republic, and 25 % at Lopé in Gabon.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDDFFD9FA306804F60BFC1E.taxon	breeding	Breeding. Female Gray-cheeked Mangabeys exhibit a bright pink genital swelling when sexually receptive. Ovarian cycleslast c. 31 days. A single young is born every 2 - 4 yearsafter a gestation of 184 - 189 days. At birth, the naked parts of the face, ears, and ischial callosities are flesh-colored. Infants are weaned at 7 - 10 months. Females are sexually mature at three years of age and males at 5 - 7 years. Interbirth intervals average ¢. 30 months, but they are shorterif the infant dies. Individuals may live for more than 20 years in captivity.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDDFFD9FA306804F60BFC1E.taxon	activity	Activity patterns. Gray-cheeked Mangabeys are diurnal and arboreal. Depending on the time of year, groups forage for 42 - 54 % of the day, travel for 20 - 28 %, rest for 10 - 28 %, and engage in social activities for 6 - 8 %.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDDFFD9FA306804F60BFC1E.taxon	biology_ecology	Movements, Home range and Social organization. Gray-cheeked Mangabeys live in multimale-multifemale groups of ¢. 20 individuals (range 6 - 30). In larger groups, there may be as many as 6 - 11 males. Studies at Mondika in the Central African Republic recorded daily movements of ¢. 1137 m. A group used 167 ha over the course of a year. Home ranges are variable in size but are generally large, 100 - 400 ha, and overlap extensively with other groups. Groups show a tendency to avoid each other. Large groups may split, and smaller subgroups can travel and forage separately for as long as three weeks at a time. In captivity, males and females form dominance hierarchies. Males usually leave their natal groups at maturity. As in other species of crested mangabeys, Gray-cheeked Mangabeys give a species-typical, low-pitched loud call (“ whoop-gobble ”), produced using an extralaryngeal resonating air sac. This loud call can be heard 500 - 1200 m away. They whoop, pause for aboutfive seconds, and then gobble for about three seconds. When they whoop, their body jerks forward slightly, lips remain closed and are pushed forward into a pout, and cheeks billow out. Theysit tense and attentive, the shoulders shake slightly, and the eyes are half closed. When gobbling, their shoulders shake up and down, their mouth is closed, and their head is slightly raised. These loud calls are given especially by high-ranking individuals. Gray-cheeked Mangabeys grunt a lot and in numerous contexts; e. g. when feeding, resting, traveling, approaching other individuals, or after copulation. Grunts vary in intensity from soft and slow to loud and rapid. Chuckles (or repeated staccato barks) are given in a variety of alarm or surprise situations such as the appearance of duikers (Cephalophus), suids such as Forest Hogs (Hylochoerus meinertzhageni) and Red River Hogs (Potamochoerus porcus), raptors, and other possible predators. Predators include the crowned hawk-eagle (Stephanoaetus coronatus) and Chimpanzees (Pan troglodytes). Population densities range from 10 ind / km? to 20 ind / km? in Cameroon and Gabon. Like other species of crested mangabey, the Gray-cheeked Mangabeyis often found in polyspecific associations with arboreal guenons (Cercopithecus), red colobus (Piliocolobus), and black-and-white Colobus (Colobus). In a number ofareas, it is sympatric with the Red-capped Mangabey (Cercocebus torquatus) or the Agile Mangabey (C. agilis).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDDFFD9FA306804F60BFC1E.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Least Concern on The [UCN Red List. The IUCN Red List recognizes the form zenkeri (not considered here) as a subspecies of the Gray-cheeked Mangabey. The Gray-cheeked Mangabeyis listed as Class B in the African Convention on the Conservation of Nature and Natural Resources. It is hunted for food in some areas and occasionally shot as a crop pest. Destruction and conversion of forest habitats also constitute threats. Gray-cheeked Mangabeys are known to occur in various protected areas: Dja Biosphere Reserve, Douala-Edéa Wildlife Reserve, and Campo-Ma’an, Boumba-Bek, Nki, and Lobéké national parks in Cameroon; Rio Campo, Monte Alen, and Los Altos de Nsork national parks in Equatorial Guinea; Loango, Lopé, and Minkebé national parks in Gabon; Nouabalé-Ndoki and Odzala national parks in the Republic of Congo; and Dzanga-Ndoki National Park and Ngotto Classified Forest Reserve in the Central African Republic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDCFFD9FAFD613BF8C4F3F4.taxon	materials_examined	Cameroon, Edea, 3 ° 30 ’ N, 10 ° 05 ’ E.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDCFFD9FAFD613BF8C4F3F4.taxon	discussion	L. osmani was first described as a subspecies. Reviewing the taxonomy of the genus in 2007, C. P. Groves considered L. osmani to be a full species. It probably hybridizes with L. albigena in the Dja Biosphere Reserve, Cameroon. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDCFFD9FAFD613BF8C4F3F4.taxon	distribution	Distribution. SE Nigeria (Gashaka-Gumti National Park) and WC Cameroon, from Edea Province and across the Sanaga River to the Cameroon Plateau. The distribution needs to be reassessed.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDCFFD9FAFD613BF8C4F3F4.taxon	description	Descriptive notes. Head — body 45 - 73 cm (males) and 50 - 56 cm (females), tail 73 — 100 cm (males) and 67 - 80 cm (females); weight 6.1 - 9 kg (males) and 4.7 - 6.5 kg (females). The Rusty-mantled Mangabey is a slender crested mangabey, with relatively long extremities and a long tail. Upper body is black with a brownish tinge, mantle is rusty-brown or tobacco-brown with the midline not much darker, and underside is yellowish-gray, with a more yellow tinge anteriorly. Arms are pale, with black hands. Crown is black, with a somewhat swept-back tuft and apparently no “ horns, ” as are found in the Gray-cheeked Mangabey (L. albigena) and Johnstons Mangabey (L. johnstoni). Cheeks are bright grayish-white or golden-white, and shoulder mantle is reddish brown.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDCFFD9FAFD613BF8C4F3F4.taxon	biology_ecology	Habitat. Primary and secondary tropical, semi-deciduous forest. The Rusty-mantled Mangabey seemsto occupy higher elevations at 600 m and higher, except in the Edea District on the coast to the north of the Sanaga River.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDCFFD9FAFD613BF8C4F3F4.taxon	food_feeding	Food and Feeding. Rusty-mantled Mangabeys eat seeds — undoubtedly an important component of the diet — along with fruits, young leaves, bark, flowers, and animal prey (invertebrates and small vertebrates). Their diets and feeding behavior have not been studied in the wild.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDCFFD9FAFD613BF8C4F3F4.taxon	breeding	Breeding. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDCFFD9FAFD613BF8C4F3F4.taxon	activity	Activity patterns. Rusty-mantled Mangabeys are diurnal and arboreal.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDCFFD9FAFD613BF8C4F3F4.taxon	biology_ecology	Movements, Home range and Social organization. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDCFFD9FAFD613BF8C4F3F4.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Least Concern on The [UCN Red List (under L. albigena). The Rusty-mantled Mangabey is listed as Class B in the African Convention on the Conservation of Nature and Natural Resources. Hunting for bushmeat and habitat loss through agricultural expansion are probably its major threats. The Rusty-mantled Mangabey is the least known of the crested mangabeys, and its distribution, ecology, and conservation status need urgent assessment. It is probably now locally extinct, for example, in Takamanda National Park in Cameroon and in the Okwango Division of Cross River National Park in Nigeria. It is known to occur in Gashaka-Gumti National Park in Nigeria and Mbam-Djerem National Park in Cameroon.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDCFFDAFAF6695BFD62F789.taxon	materials_examined	Congo-Zaire, near Lake Tanganyika. Restricted by Lorenz von Liburnau in 1917 to Semliki or Ituri Forest, and the type locality was fixed by E. Schwarz in 1910 to (present-day) Burundi, north-east of Lake Tanganyika.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDCFFDAFAF6695BFD62F789.taxon	discussion	L. johnston: was first described as a subspecies of L. albigena, but C. P. Groves, in reviewing the taxonomy of the genus in 2007, considered it to be a full species. He also separated part of the L. johnstoni population as L. ugandae. Whether L. johnston: intergrades with L. ugandae is not known. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDCFFDAFAF6695BFD62F789.taxon	distribution	Distribution. DR Congo (E and S of the Ubangi River and N of the Congo River), W Rwanda, and W Burundi; possibly in NW Uganda (W of the Albert Nile) and in the Republic of South Sudan.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDCFFDAFAF6695BFD62F789.taxon	description	Descriptive notes. Head-body 45 - 73 cm (males) and 50 - 56 cm (females), tail 73 - 100 cm (males) and 67 - 80 cm (females); weight 6.1 - 9 kg (males) and 4.7 - 6.5 kg (females). Johnston's Mangabey is a slender crested mangabey (Lophocebus), with relatively long extremities and a long tail. Fur is darkish brown to black, dorsally and ventrally. Mantle is a slightly lighter, gray-brown. Crown hairs are swept back, with long eyebrow tufts (“ horns ”), and cheeks are light gray-brown passing to white inferiorly, but very thinly haired. Albinism is common.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDCFFDAFAF6695BFD62F789.taxon	biology_ecology	Habitat. Primary and secondary tropical, semi-deciduous forest.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDCFFDAFAF6695BFD62F789.taxon	food_feeding	Food and Feeding. The diet ofJohnston’s Mangabey is undoubtedly similar to other crested mangabeys and probably includes seeds as a major component, along with fruits, young leaves, bark, flowers, and animal prey (invertebrates and small vertebrates).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDCFFDAFAF6695BFD62F789.taxon	breeding	Breeding. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDCFFDAFAF6695BFD62F789.taxon	activity	Activity patterns. Johnston’s Mangabeys are diurnal and arboreal.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDCFFDAFAF6695BFD62F789.taxon	biology_ecology	Movements, Home range and Social organization. In the Ituri Forest in the DR Congo, group size is 6 - 14 individuals, with an abundance estimate of 6 - 9 ind / km. In lowland areas of Kahuzi-Biéga National Park, they are more abundant, estimated at 15 - 9 ind / km with an average group size of 21 - 4 individuals. In most of its distribution, it is sympatric with the Agile Mangabey (Cercocebus agilis) and is often found in polyspecific associations with colobines and arboreal guenons.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDCFFDAFAF6695BFD62F789.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Least Concern on The [UCN Red List (under L. albigena). Johnston’s Mangabey is listed as Class B in the African Convention on the Conservation of Nature and Natural Resources. It is hunted for food in some areas and occasionally shot as a crop pest. It has become scarce in several protected areas because of the civil war and rebel and military occupation ofits forests. Johnstons Mangabey is known to occur in a number of protected areas: Kibira National Park in Burundi; Bomu Wildlife Reserve, Yangambi Reserve, Maiko National Park, [turi Forest, Okapi Faunal Reserve, Kahuzi-Biéga National Park and Virunga National Park in the DR Congo; Nyungwe Forest National Park in Rwanda; and probably Ajai Game Reserve in Uganda. A reassessment of the distribution and conservation status ofJohnston’s Mangabey is urgently needed.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDFFFDAFF2C6DAEF788F69D.taxon	materials_examined	Uganda, Chagwe, Nile mouth at Lake Albert. Restricted by C. P. Groves in 2007 to Chagwe, “ a large area north of Lake Victoria, east of Kampala, west of the Nile and Jinja, and a little northwards toward Bugerere, more correctly called Kyagwe. ”	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDFFFDAFF2C6DAEF788F69D.taxon	discussion	Some experts consider L. ugandae to be a subspecies of L. albigena. It was separated from L. johnstoni by Groves in his 2007 review of the genus. Whether L. ugandae intergrades with L. johnstoni is not known. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDFFFDAFF2C6DAEF788F69D.taxon	distribution	Distribution. Uganda, along the N & NW shores of Lake Victoria, including Mabira Forest (most likely the type locality), Bujuko and Bukasa Forests, and along the E side of the Albertine Rift, especially in Kibale Forest National Park and the Bukoma area; it is also found in Sango Bay Forest and in Minziro Forest, NW Tanzania, both parts of the Minziro-Sango Bay Transboundary Site.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDFFFDAFF2C6DAEF788F69D.taxon	description	Descriptive notes. Head-body 54 - 70 cm (males) and 42 - 61 cm (females), tail 65 — 75 cm (males) and 55 - 65 cm (females); weight 6 - 10 kg (males) and 4 - 6 kg (females). As in other species of crested mangabeys (Lophocebus), fur of the Ugandan Crested Mangabey is soft and long, being generally dark gray-brown or blackish-brown with a ragged tail and a tufted crown. Eyebrow tufts form “ horns ” as they do in the Graycheeked Mangabey (L. albigena) and Johnston’s Mangabey (L. johnstoni). A longer, paler mantle on the shoulders (more prominent in the males) runs continuous with rather thin, grayish, or pale whitish cheek whiskers. Skin of the face is entirely black. Pelage characteristics of the Ugandan Crested Mangabey need to be restudied, but the evidence to date indicates that its mantle contrasts more with the general body color than it does in the pelage ofJohnston’s Mangabey. When standing quadrupedally, tail is often held vertically with the tip curved forward. The Ugandan Crested Mangabeyis probably the smallest species of the crested mangabeys. Males are larger than females, but sexual dimorphism is not as pronounced as in the other species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDFFFDAFF2C6DAEF788F69D.taxon	biology_ecology	Habitat. Primary and some secondary tropical, semi-deciduous, flooded, and gallery forest. Ugandan Crested Mangabeys are most typically found in dense evergreen forest where it prefers the middle and upperlayers of the canopy.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDFFFDAFF2C6DAEF788F69D.taxon	food_feeding	Food and Feeding. In two studies conducted in Kibale Forest National Park, diets of Ugandan Crested Mangabeys consisted of 35 - 78 % fruit, 2 - 10 % leaves, 1 - 8 % flowers, and 0 - 9 % animal matter. They used 51 and 65 different plant species, respectively. Hard foods make up a considerable proportion of the diet of all mangabey species, including the Ugandan Crested Mangabey, and their thick enamelis regarded as an adaptation.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDFFFDAFF2C6DAEF788F69D.taxon	breeding	Breeding. In Kibale Forest, the interbirth interval is ¢. 33 months, the mean length of the ovarian cycle is 47 days (range 8 - 184), and the gestation period is ¢. 186 days. In captivity, the average cycle length (30 days) and gestation (175 days) are shorter. Female Ugandan Crested Mangabeys develop a sexual swelling similar to that of female baboons. Males and females form consortships (they stay close together) during the female’s periovulatory period, but males, it seems, are unable to monopolize females at this time to the same extent as observed in some baboon species. Females typically solicit copulation from multiple males. Females vocalize during and shortly after copulation. Males produce a distinctive loud call, the “ whoop-gobble. ”	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDFFFDAFF2C6DAEF788F69D.taxon	activity	Activity patterns. Ugandan Crested Mangabeys are diurnal and arboreal. They rarely descend to the ground.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDFFFDAFF2C6DAEF788F69D.taxon	biology_ecology	Movements, Home range and Social organization. Ugandan Crested Mangabeys live in multimale — multifemale groups of five to as many as 30 individuals, but usually less than 20. Larger groups contain more than one adult male (ratio of males to femalesis 1: 2 - 4). Average home range in Kibale Forest is 155 ha, and daily movements are 0 - 8 - 1 - 5 km. Population densities are variable, from less than 4 ind / km? to 8 - 10 ind / km?. In Kibale Forest, their numbers are high at 34 ind / km?, but in gallery forest, densities can reach 77 ind / km?. Males and females probably form dominance hierarchies, but this has been confirmed only for captive animals. Males usually leave their natal groups after reaching maturity. Like other species of crested mangabey, the Ugandan Crested Mangabey is often found in polyspecific associations with Blue Monkeys (Cercopithecus mitis), Red-tailed Monkeys (C. ascanius), Ashy Red Colobus (Piliocolobus tephrosceles), and Guerezas (Colobus guereza).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDFFFDAFF2C6DAEF788F69D.taxon	conservation	Status and Conservation. CITES Appendix II. The Ugandan Crested Mangabey has not been assessed on The IUCN Red List. It is listed as Class B in the African Convention on the Conservation of Nature and Natural Resources. Ugandan Crested Mangabeys are hunted for food in some areas and occasionally shot as a crop pest. They occur in four protected areas: Kibale Forest National Park, Mabira Forest Reserve, and Bugoma Forest Reserve in Uganda and Minziro Forest Reserve in Tanzania. In Kibale Forest, the population seems to have been stable in recent decades.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDFFFDBFA376CBAFEE2F713.taxon	materials_examined	Congo-Zaire, Stanley Falls.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDFFFDBFA376CBAFEE2F713.taxon	discussion	The population of L. aterrimus in the south-western part of their distribution was described as the subspecies opdenboschi, but following C. P. Grove’s review in 2007, itis regarded here to be a full species (L. opdenboschi). L. aterrimus replaces L. johnston: south of the Congo River. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDFFFDBFA376CBAFEE2F713.taxon	distribution	Distribution. Endemic to DR Congo, in the C Congo Basin S & W of the Congo-Lualaba River System.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDFFFDBFA376CBAFEE2F713.taxon	description	Descriptive notes. Head-body 40 - 62 cm, tail 55 - 85 cm; weight 8 kg (males) and 4 - 5 — 6 - 7 kg (females). The Northern Black Crested Mangabey is a slender mangabey, with relatively long extremities and tail, and typified by its coarse and almost entirely black color. It has a high conical crest that rises to a point at the back of the head, but no brow fringe or shoulder mantle. Its face is naked and slate-black with orangey-red eyes. Cheek whiskers are sepia-gray (contrasting with the black of the body) and notably thick and elongated; they sweep back and up with a slight outward and forward curve. Female Northern Black Crested Mangabeys are smaller than males. Albinism and leucism (reduced pigmentation) are fairly common.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDFFFDBFA376CBAFEE2F713.taxon	biology_ecology	Habitat. A variety of forest types including semi-deciduous, gallery, and swamp forests. In Salonga National Park, the Northern Black Crested Mangabey occurs in swamp forests, but it does not occupy swamp in Lomako. It uses all forest levels (especially the middle canopy) and seldom descends to the ground.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDFFFDBFA376CBAFEE2F713.taxon	food_feeding	Food and Feeding. The diet of the Northern Black Crested Mangabey contains mainly fruits (as much as 57 % of the diet) and seeds (up to ¢. 30 % of the diet). It also eats young leaves (up to 16 % of the diet), flowers and nectar (up to 17 - 5 %), bark, gums, and animal prey (including insects and small reptiles). In Salonga National Park, Northern Black Crested Mangabeys show high rates of nectarivory in some months.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDFFFDBFA376CBAFEE2F713.taxon	breeding	Breeding. Female Northern Black Crested Mangabeys exhibit a pink genital swelling when sexually receptive. In captivity, the menstrual cycle is 21 - 40 days (average 26 days).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDFFFDBFA376CBAFEE2F713.taxon	activity	Activity patterns. Northern Black Crested Mangabeys are diurnal and arboreal. An activity budget recorded at La Tumba in the DR Congo was 36 - 2 % foraging and traveling, 38 - 3 % resting (especially in late morning and late afternoon), and 24 - 0 % engaging in social behavior; very little time was spent exclusively traveling (1 - 5 %). They use all levels of the forest canopy, to the tallest canopy-emergents, but lower and middle canopies (12 - 30 m above ground) are used most for foraging and traveling. They rest mostly at 24 — 42 m.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDFFFDBFA376CBAFEE2F713.taxon	biology_ecology	Movements, Home range and Social organization. The Northern Black Crested Mangabey lives in multimale — multifemale groups of 11 - 19 individuals. The home range of a group of 19 individuals (with three adult males five adult females) at Lake Tumba (DR Congo) was 70 ha, and that of another group of the same size was 48 ha. The groups showed noterritorial defense, and home ranges overlapped by 60 - 75 %. Males disperse. A population density of 69 ind / km * has been recorded at Lake Tumba and 73 ind / km? at Lomako. Primary vocalizations of the Northern Black Crested Mangabey are very similar to those of the closely related Gray-cheeked Mangabey (L. albigena). “ Whoop-gobble ” calling recorded for the Northern Black Crested Mangabey indicate that it is a male call (as has been found for the Gray-cheeked Mangabey) and may function in intergroup location and spacing and in male dominance (males within the group and between males in different groups). About 70 % of all whoop-gobbles are heard in the morning from sunrise to ¢. 11: 00 h. Grunts are given predominantly in agonistic situations between males. Staccato barks of the Northern Black Crested Mangabey (resembling distant braying of a donkey) are alarms, given when seeing humans and their main predators such as crowned hawk-eagles (Stephanoaetus coronatus). There is also evidence that Bonobos (Pan paniscus) prey on Northern Black Crested Mangabeys. They frequently associate with other species of arboreal monkeys, and alarm calls of Red-tailed Monkeys (Cercopithecus ascanius) and Wolf's Monkeys (C. wolf) that travel with them elicit their staccato barks.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDFFFDBFA376CBAFEE2F713.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Near Threatened on The IUCN Red List. The Northern Black Crested Mangabey is listed as Class B in the African Convention on the Conservation of Nature and Natural Resources. It is hunted for food and may be threatened by loss of habitat from logging. Little is known about its status. It is subject, however, to intensive, uncontrolled hunting for its meat in most parts of its distribution, and it is also vulnerable to loss of forest habitat. The Northern Black Crested Mangabey occurs in Salonga National Park, Tumba-Lediima Reserve, and Lomako-Yokokala Faunal Reserve in the DR Congo.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDEFFDBFFF66C27F6F5FCC1.taxon	materials_examined	Congo-Zaire, Mwilambongo (= Mwiliambongo), 4 ° 53 ’ S, 19 ° 42 ’ E.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDEFFDBFFF66C27F6F5FCC1.taxon	discussion	L. opdenboschi was formerly considered a subspecies of L. aterrimus. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDEFFDBFFF66C27F6F5FCC1.taxon	distribution	Distribution. SW DR Congo (forests along the Kwilu, Wamba, and Kwango rivers) and N Angola.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDEFFDBFFF66C27F6F5FCC1.taxon	description	Descriptive notes. Head-body 40 - 62 cm, tail 55 - 85 cm; weight 8 kg (males) and 4 - 5 — 6 - 7 kg (females). The Southern Black Crested Mangabey is very similar to the Northern Black Crested Mangabey (L. aterrimus), but it is entirely black and has longer, laxer fur. Cheek whiskers are not sepiagray, as in the Northern Black Crested Mangabey, but black without a noticeable curve. Crown crest of the Southern Black Crested Mangabey is more flat and not pyramidal; it is also broader than in the Northern Black Crested Mangabey and slopes back and slightly up. Albinism has been reported in the Southern Black Crested Mangabey. Females are smaller than males.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDEFFDBFFF66C27F6F5FCC1.taxon	biology_ecology	Habitat. Primary and some secondary rainforest, and swamp and gallery forest.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDEFFDBFFF66C27F6F5FCC1.taxon	food_feeding	Food and Feeding. Diets of Southern Black Crested Mangabeys are undoubtedly similar to other crested mangabeys and probably include seeds as a major component, along with fruits, young leaves, bark, flowers, and animal prey (invertebrates and small vertebrates).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDEFFDBFFF66C27F6F5FCC1.taxon	breeding	Breeding. There is no specific information available for this species, but female Southern Black Crested Mangabeys exhibit a pink genital swelling when sexually receptive.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDEFFDBFFF66C27F6F5FCC1.taxon	activity	Activity patterns. Southern Black Crested Mangabeys are diurnal and arboreal.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDEFFDBFFF66C27F6F5FCC1.taxon	biology_ecology	Movements, Home range and Social organization. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDEFFDBFFF66C27F6F5FCC1.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Data Deficient on The [UCN Red List (as L. aterrimus opdenboschi). The Southern Black Crested Mangabey is listed as Class B in the African Convention on the Conservation of Nature and Natural Resources. It is a poorly known, with no recent information on its population status or limits of its distribution. Mostlikely, it is hunted for food and may be threatened by loss of habitat because of logging. The Southern Black Crested Mangabey is not known to occur in any protected areas.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDEFFE4FAFE615BF92BF70E.taxon	materials_examined	Rungwe-Livingstone (09 ° 07 ° S to 09 ° 11 ° S and 33 ° 40 ° E to 33 ° 55 FE), Southern Highlands, Tanzania.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDEFFE4FAFE615BF92BF70E.taxon	discussion	R. kipunji was originally allocated to the genus Lophocebus based on its non-contrasting black eyelids and arboreal nature, but later placed in a new monotypic genus Rungwecebus on the basis of molecular and morphological data. Rungwecebus was initially questioned, but additional molecular evidence and morphometric analyses support its phylogenetic position and taxonomic status. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDEFFE4FAFE615BF92BF70E.taxon	distribution	Distribution. Endemic to S Tanzania in just two isolated sites, the Southern Highlands and the Udzungwa Mts.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDEFFE4FAFE615BF92BF70E.taxon	description	Descriptive notes. No exact adult measurements are available, but the head-body length is estimated at ¢. 85 - 90 cm, the tail a little longer; weight c ¢. 10 - 15 kg. The Kipunji is a mostly brown, medium-sized, long-tailed, arboreal monkey. Sexes are similar in color, and adult females are estimated to be 90 % the weight of adult males. Facial skin, including eyelids, is black. Muzzle is bare, relatively elongated, and black. Suborbital fossa is pronounced. Eyes are brown. Cheek-whiskers are long, extending laterally and curving downward. Crown has a prominent, long, broad, and erect crest of dark grayish-brown hair. Shoulder mane (mantle or cape) of adult males, variable in color and length, typically has long, straight, cinnamon-brown hairs. Lower dorsum is light to medium grayish-brown. Forelimbs are a dark grayish-brown. Upper hindlimbs are dark rufous-brown, and lower hindlimbs are cinnamon. Hands and feet are black. Ventrum is off-white. There is a patch on the upper chest that is close to cinnamon. Individual hairs are long and straight, without banding or speckling. Ischial callosities are pink and fused in males and not fused in females. Tail is dusky-brown over the proximal half and off-white with interspersed darker hairs over the distal half. Its pelage is smooth, not shaggy or lax. Tail is carried loosely and parallel to, or below, the plane of the back, curving downward to the level of the feet during locomotion. It is not held arched over the back when standing, nor held vertically, except as a semi-prehensile support. Infants are less hirsute with a reddish tinge to their pelage.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDEFFE4FAFE615BF92BF70E.taxon	biology_ecology	Habitat. Degraded montane and upper montane forest at 1500 - 2450 m above sea level in Rungwe-Kitulo to pristine submontane forest at 1300 - 1750 m in Ndundulu. In Rungwe-Kitulo, the canopy is often broken at 10 - 30 m with emergents as high as 35 m. Kipunjis prefer steep-sided gullies and valley edges. Ridges and open areas are usually avoided. Numbers of Albizia gummifera (Fabaceae), Casearia battiscombei (Salicaceae), Entandrophragma sp. (Meliaceae), Macaranga kilimandscharica (Euphorbiaceae), Ocotea usambarensis (Lauraceae), Olea capensis (Oleaceae), Parinari excelsa (Chrysobalanaceae), Podocarpus latifolius (Podocarpaceae), and Prunus africana (Rosaceae) have been greatly reduced by logging. Thick undergrowth is typical, with tree ferns Cyathea manniana (Cyatheaceae), wild bananas Ensete ventricosum (Musaceae), and large stands of bamboo Sinarundinaria alpina (Poaceae), common in the south and south-east of Mount Rungwe and in the north-west of Livingstone. Kipunjis rarely frequent bamboo, and they leave the forest only to raid nearby crops, especially in February — April. In areas inhabited by the Kipunji, annual rainfall is ¢. 1850 - 2500 mm. Mean annual rainfall from 1968 to 2008 was 2133 mm. The wet season is in November — May, and the dry (colder) season is in June-October. In Ndundulu, the primary forest, dominated by P. excelsa, is mostly undisturbed and its canopy unbroken, reaching heights of 40 - 50 m.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDEFFE4FAFE615BF92BF70E.taxon	food_feeding	Food and Feeding. The Kipunji is omnivorous. During 9498 hours of observation of 34 groups in the Southern Highlands, 122 food plants from 60 families were recorded in the diet (64 tree species, 30 herbs, nine climbers, seven shrubs, six lianas, three grasses, and three ferns). The diet comprises mature leaves (22 %), unripe fruits (14 %), ripe fruits (13 %), young leaves (12 %), bark (11 %), stalks (9 %), flowers (9 %), pith (7 %), insects (2 %), and seed pods, rhizomes, shoots, tubers, epiphytes, ground herbs, climbers, moss, fungi and lichen (2 %). Kipunji ate soil on two occasions. They also raid maize, bean, pea, sweet potato, and banana crops and granaries if close to the forest edge. The most frequently eaten plant is Macaranga capensis (Euphorbiaceae). Other commonly eaten plants include Ilex mitis (Aquifoliaceae), Psydrax parviflora (Rubiaceae), Chrysophyllum gorungosanum (Sapotaceae), Bersama abyssinica (Melianthaceae), Myrianthus holstii (Cecropiaceae), Tabernaemontana stapfiana and Landolphia buchananii (both Apocynaceae), Urera hypselodendron (Urticaceae), Tarenna pavettoides and Multidentia crassa (both Rubiaceae), Cassipourea gummiflua (Rhizophoraceae), Allophylus abyssinicus (Sapindaceae), Parinari excelsa, Entandrophragma excelsum (Meliaceae), Ficus thonningii (Moraceae) and Polyscias fulva (Araliaceae). They are able to open large, hard fruits and break apart dead wood in search of grubs. The Kipunji appears to be more folivorous during the dry season and more frugivorous during the wet season. Most foraging occurs in the early morning and late afternoon.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDEFFE4FAFE615BF92BF70E.taxon	breeding	Breeding. Little is known about breeding in the Kipunji. In Rungwe-Kitulo, Kipunjis were seen mating on four occasions, all in late September. One pair moved ¢. 200 m from the main group, copulated twice in ten minutes and again 45 minutes after that. Copulations lasted 3 - 7 seconds. Infants are carried beneath the female. Instances of “ kidnapping ” are accompanied by loud screams and chasing. The female’s genitals swell during the periovulatory period. Single offspring are born; twins have not been recorded. Infants are born throughout the year, and there is no evidence of a birth season or birth peak. The gestation period is 6 - 7 months.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDEFFE4FAFE615BF92BF70E.taxon	activity	Activity patterns. Kipunjis are diurnal and arboreal, foraging and traveling mainly at 10 m or more above the ground. It occasionally feeds on the forest floor, particularly in the dry season. The Kipunji also goes to the ground to cross degraded forest patches and to avoid intragroup conflict and predators. Kipunjis are tolerant of low temperatures; in parts of Rungwe-Kitulo, temperatures can drop to - 3 ° C in June-August. They sleep high up, usually above 20 m, in branches oftrees such as B. abyssinica, M. holstii, Maesa lanceolata (Myrsinaceae), and Syzygium guineense (Myrtaceae). They usually rest for 30 - 60 minutes in the middle of the day. They are most active at 08: 00 - 11: 30 h and 15: 00 - 18: 00 h.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDEFFE4FAFE615BF92BF70E.taxon	biology_ecology	Movements, Home range and Social organization. Groups of Kipunjis in Rungwe-Kitulo travel a mean of 1 - 3 km / day (range 0 - 99 - 17, SE = 0 - 15 km, n = 235 days), and group members usually remain within 300 m of each other, except when disturbed. Mean home range in Mount Rungwe is 306 ha (116 - 430 ha, SE = 67 ha, n = 4 groups). The Kipunji is social and notterritorial. Groups comprise several adult males and several adult females. Group sizes are 20 - 39 individuals in Rungwe-Kitulo and 15 - 25 in Ndundulu. In 34 groups that were closely monitored in Rungwe-Kitulo, there were up to five clinging infants per group. There is no evidence ofsolitary individuals. In Rungwe-Kitulo, groups of Kipunjis form polyspecific associations with the Angolan Colobus (Colobus angolensis) and Sykes’s Monkey (Cercopithecus albogularis moloneyi), especially in early morning and late afternoon. The three species often sleep in neighboring trees. Groups of Kipunji at Ndundulu form polyspecific associations with Angolan Colobus, Sykes’s Monkeys, and Udzungwa Red Colobus (Piliocolobus gordonorum). Grooming is observed most often in the early afternoon. Receptive females are often groomed by young males, although female — female, female-male, and adult — juvenile grooming are also observed. Intergroup territorial displays occur when ranges overlap. Disputes are often highly vocal in Rungwe-Kitulo, usually involving adult males, but sometimes females and juveniles also call. No intergroup physical contact was observed in over 772 hours of direct observation. In Rungwe-Kitulo, adult males emit a distinctive, loud, low-pitched “ honk-bark, ” most evident when conspecific groups meet and when threatened or disturbed. Adult females also give this call but less often than males. Honk-barks can be heard by the human ear up to 1 km away and are used as a groupspacing mechanism. The honk-bark is qualitatively and quantifiably different from the “ whoop-gobble ” of species of crested mangabeys (Lophocebus) and has some structural congruence to the “ roar-grunt ” of baboons (Papio). At least ten other call types of Kipunjis have been identified, including contact grunts. High-pitched, sharp screams are given in alarm when crowned hawk-eagles (Stephanoatus coronatus) are seen and during intragroup agonistic encounters.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFDEFFE4FAFE615BF92BF70E.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Critically Endangered on The IUCN Red List. The Kipunji is known from just two sites of largely non-forested land separated by ¢. 350 km. One fragmented population occurs at 1750 - 2450 m above sea level in only 42 km * of Rungwe-Kitulo Forest in the Southern Highlands. This forest includes Mount Rungwe Nature Reserve of 150 km ” and the Livingstone Forest of 191 km? (in Kitulo National Park of 412 km?). Mount Rungwe Nature Reserve and Livingstone Forest are connected by the Bujingijila Corridor, a narrow (less than 2 km wide) strip of degraded forest. The Kipunji inhabits the wetter forest of southern Mount Rungwe, and isolated groups are scattered in the northern and southern areas of Livingstone Forest. The extent of their occurrence in Mount Rungwe and Livingstone is 23 - 7 km * and 18 - 3 km?, respectively. Anecdotal evidence points to the historic presence of Kipunjis in other forests of the Southern Highlands. The second population of the Kipunji occurs at elevations of 1300 - 1750 m in the Vikongwa Valley, Ndundulu Forest, in the Kilombero Nature Reserve in the Udzungwa Mountains. The known extent of occurrence in the Ndundulu Forest is at least 5 - 3 km?, but is likely to be much larger. The total population of the Kipunji is estimated to be 1117 animals in 38 groups. Of these, 1042 individuals occur in Rungwe-Kitulo in 34 groups, and there are only 75 individuals in Ndundulu in four groups. Kipunjis are Africa’s most threatened monkey and one of the world’s most threatened primates. Threats to Kipunjis are considerable. The Rungwe-Kitulo forests are severely degraded; logging, charcoalmaking, illegal hunting, and unmanaged resource extraction are common. The narrow Bujingijila Corridor linking Mount Rungwe to Livingstone Forest and corridors joining the northern and southern sections of Livingstone have been encroached upon and degraded by farmers. Without immediate conservation intervention, these forests will be fragmented, resulting in the isolation of Kipunji subpopulations, some of which are unlikely to be viable over the long term. Protecting these corridors is of the highest priority for the conservation of the Kipunji. It is hunted by local people, especially on farms. Conflicts with local famers become more common as forest disturbance and loss increase. Mount Rungwe was raised to the status of a Nature Reserve in 2009, but it remains largely unmanaged. The smaller population of Kipunjis in Livingstone Forest is within Kitulo National Park, but swift and effective conservation action 1 s needed in both areas. Although Ndundulu Forest is in excellent condition, and largely undisturbed, Kipunjis are present in low numbers, and reasons for this are unclear. Kilombero Nature Reserve was established in 2007, incorporating Ndundulu, but as with Mount Rungwe, there is as yetlittle conservation management. The Kipunji is serving as a “ flagship ” genus and species for the conservation of the Southern Highlands and Udzungwa Mountains. Their presence at both sites demonstrates that the Southern Highlands and Eastern Arcs were linked and are zoologically even more aligned than had previously been thought.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE3FFE6FF2C64A3F818F50C.taxon	materials_examined	Africa. Restricted by J. A. Allen in 1925 to “ Kenya, inland from Mombasa. ”	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE3FFE6FF2C64A3F818F50C.taxon	discussion	P. cynocephalus hybridizes with P. anubis in the eastern part of Tsavo and Amboseli national parks in Kenya. There is a broad clinal hybrid zone of PF. cynocephalus x P. anubis between Laikipia district, just to the north-east and east of Mount Kenya, and the Lower Tana River, Kenya coast. Baboons in this more than 200 km-wide region are intermediate and are difficult to allocate to either P. cynocephalus or P. anubis (baboons become increasingly “ yellow-like ” in their phenotypes toward the Kenya coast). Whether their distribution overlaps with P. hamadryas in Somalia is not clear. In respective contact zones in Zambia, they hybridize with P. kindae and P. ursinus griseipes. Molecular studies also indicate that P. cynocephalus hybridized with the Kipunji (Rungwecebus kipunji) in southern Tanzania. Historically, several forms of P. cynocephalus have been described, of which a few might merit subspecies status (e. g. Luangwa Valley Yellow or Dwarf Chacma Baboon, P. c. jubilaeus). P. c. kindae is recognized here as a separate species. Limits to distributions of the two subspecies are poorly known. Two subspecies recognized.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE3FFE6FF2C64A3F818F50C.taxon	distribution	Subspecies and Distribution. P. c. cynocephalusLinnaeus, 1766 — Tanzania, Malawi, EZambia (EofLuangwaRiver), andNMozambique. P. c. ibeanus Thomas, 1893 — SE Somalia, E Kenya (including Manda I), and NE Tanzania; possibly in extreme SE Ethiopia but the N limit in Somalia is not known.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE3FFE6FF2C64A3F818F50C.taxon	description	Descriptive notes. Head-body 62 - 85 cm (males) and 51 - 69 cm (females), tail 53 — 66 cm (males) and 34 - 57 cm (females); weight 20 - 28 kg (males) and 8 - 13 kg (females). Sexual dimorphism in body weights of Yellow Baboons is pronounced, with female body mass only 54 - 56 % of male body mass. The Yellow Baboon differs from Chacma (P. ursinus) and Olive (P. anubis) baboons by its relatively lanky form, smaller head, and elongated, slender limbs. Upper body of the Yellow Baboon is yellowishfawn to yellowish-gray, with white on the underside, inner surfaces of limbs, cheeks, sides of the muzzle, and fringes of hands and feet. Males have a slight median nuchal crest of long hair, but little or no shoulder mane. Flank hairs on both sexes are elongated and form an inconspicuous fringe. Face, palms of hands, and soles of feet are black. Nostrils do not protrude beyond the upperlip. Ischial callosities are small and grayish-black; in males, they form a continuous ridge but in females they are separated by the genitalia. Tail is often carried almost vertically, then falling downward in a low, “ broken ” fashion. Females show prominent swellings of their perineum when sexually receptive. Infants are born with black natal coats that change to adult coloration at 3 - 6 months of age. Subspecific morphological differences are slight among Yellow Baboons. In the “ Central Yellow Baboon ” (P. ¢. cynocephalus), furis straight, and males lack any trace of a shoulder mane. In the “ Northern Yellow Baboon ” (P. c. ibeanus), fur is wavy, and males have a trace of a shoulder mane.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE3FFE6FF2C64A3F818F50C.taxon	biology_ecology	Habitat. Largely fire-climax miombo (Brachystegia, Fabaceae) woodland. In this zone and especially to the north-east, Yellow Baboons occupy dry bushland, thickets, steppes, and the coastal littoral (including mangroves) zone, as long as access to fresh water sources for drinking is provided. Yellow Baboons are able to persist in secondary and highly fragmented vegetation, and they can adapt to disturbed habitats near human settlements, living in cultivated areas if there is native habitat for refuge.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE3FFE6FF2C64A3F818F50C.taxon	food_feeding	Food and Feeding. Like all baboons, Yellow Baboons are opportunistic omnivores. In parts of their distribution, they feed primarily on seeds, fleshy, pods, and exudates of leguminous trees. It has been hypothesized that they prefer foods with an unusual chemistry, implying that they have special digestive adaptations, possibly explaining why the boundaries of its distribution do not follow any geographic discontinuities but coincide very closely with the distribution of a specific plant community. Nevertheless, in other areas of their distribution, Yellow Baboons have diets very similar to those of other baboons. They feed on grasses, shoots, fungi, lichens, and animal prey (including invertebrates, lizards, birds, and small mammals).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE3FFE6FF2C64A3F818F50C.taxon	breeding	Breeding. In Amboseli, female Yellow Baboons show theirfirst sexual swellings at 4 - 6 years of age and have their first infant, on average, at 5 - 9 years. Their menstrual cycle takes 26 - 52 days. Male maturation takes longer, and they are considered adults at c. 8 years old. The gestation period of the Yellow Baboon is ¢. 180 days. Births occur throughout the year, and interbirth intervals are usually 22 - 27 months. Males try to monopolize access to females that are sexually receptive. A male and a female form a temporary consortship (several minutes to days). Most mating happens during these consort periods. The consorting male is often challenged by other males, and male consort partners may change several times during any one receptive period. Hence, female Yellow Baboons usually mate with several partners. Females give copulation calls, i. e. they vocalize during and shortly after copulation. Life expectancy in wild females is c. 14 - 15 years, but females have been recorded living up to 27 years in the wild. Because of the social system and dispersal pattern of the Yellow Baboon, itis more difficult to estimate male life span in the wild.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE3FFE6FF2C64A3F818F50C.taxon	activity	Activity patterns. Yellow Baboon are diurnal and mainly terrestrial.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE3FFE6FF2C64A3F818F50C.taxon	biology_ecology	Movements, Home range and Social organization. Home range size and daily movement vary considerably depending on habitat, group size, and season. If food is abundant, daily movements are less than 4 km, but in the dry season, Yellow Baboons can move 8 - 10 km / day. Besides food, home ranges contain essential localized resources such as water and sleeping sites. Yellow Baboons usually sleep in trees. Like Chacma and Olive baboons, they live in multimale-multifemale groups of 20 - 80 individuals. Groups are rarely less than 20 or more than 100 individuals. Adult sex ratios within groups can vary substantially, but there are typically more females than males. Groups are not substructured into one-male units, as in Hamadryas Baboons (P. hamadryas) or Geladas (Theropithecus gelada), although solitary one-male units have been observed. Most male Yellow Baboons emigrate from their natal groups at 7 - 13 years of age. Secondary emigration into a third group has also been recorded. Females remain in their natal groups and form linear dominance hierarchies. High ranking mothers support their daughters during rank conflicts, resulting in a dominance rank of the daughter just below the mother. Rank positions are “ heritable, ” and dominance relationships among matrilines within a group can remain stable over generations. Males also establish a dominance hierarchy that usually regulates access to receptive females. The dominant male in a group can sire more than 80 % of all offspring, but low-ranking males can form coalitions to challenge higher-ranking males and to take over sexually receptive females. The social network among related females is the most stable social structure within a group of Yellow Baboons. Grooming and otheraffiliative behavior are exchanged predominantly within these networks. Studies suggest that these tight relationships provide a fitness advantage for females. Females with a tighter network live longer and reproduce more successfully. Females and males may also form bonds outside sexual consortships, called “ friendships. ” These relationships likely benefit a female via protection for herself and her offspring against sexual harassment and infanticide by other males. The male benefits because it may protect its own offspring or it may invest in future sexual relationship with respective females.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE3FFE6FF2C64A3F818F50C.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Least Concern on The IUCN Red List, including both subspecies. Yellow Baboons are listed as vermin in the African Convention on the Conservation of Nature and Natural Resources. They are used extensively in biomedical research and often shot as crop pests. They are widespread and locally common, but patchily distributed over their extensive distribution. They have been locally displaced by agriculture, development, and infrastructure projects. The Northern Yellow Baboon occurs in a number of protected areas, including Amboseli National Park and Tana River National Primate Reserve in Kenya and Mikumi, Saadani, Katavi and possibly Ruaha national parks in Tanzania. Its status in Somalia and Ethiopia is unclear. The Central Yellow Baboon is also widespread and common and occurs in various protected areas: Ruaha National Park (possibly), Udzungwa Mountains National Park, and Selous Game Reserve in Tanzania; Niassa Reserve, Quirimbas National Park and Gilé National Reserve in Mozambique; and Kasungu and Liwonde national parks in Malawi. There are no major range-wide threats believed to be resulting in any significant population decline of the Central Yellow Baboon.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE3FFE7FA346E24FB0EF786.taxon	materials_examined	Congo-Zaire, Lulua District, Kinda.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE3FFE7FA346E24FB0EF786.taxon	discussion	P. kindae has previously been considered a subspecies of P. cynocephalus. It is recognized here as a full species because it is as morphologically and genetically distinctive as the five other baboon species. Furthermore, preliminary behavioral observations suggest significant differences compared with P. cynocephalus. This species hybridizes with P. wrsinus griseipes and P. c. cynocephalus in respective contact zones in Zambia.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE3FFE7FA346E24FB0EF786.taxon	distribution	Distribution. Extreme SW Tanzania (possibly N to Mahale Mountains National Park), S DR Congo, W Zambia (W of Luangwa River), and N Angola.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE3FFE7FA346E24FB0EF786.taxon	description	Descriptive notes. Head-body ¢. 58 cm (males) and c ¢. 56 cm (females), tail c. 53 cm (males) and c. 47 cm (females); weight c. 16 kg (males) and c. 10 kg (females). The Kinda Baboon is smaller and more gracile and slender than other baboons. It has a very short muzzle. It is the least sexually dimorphic in size; female body mass is 63 % of male body mass. The adult male Kinda Baboon has a mass matching that of adult female Yellow (P. eynocephalus) and Chacma (P. ursinus) baboons. Fur of the Kinda Baboon is yellowish brown dorsally, and paler and creamy ventrally. Pelage has a soft, silky texture, distinctive from Yellow Baboons. The Kinda Baboon has a pronounced crest of hair on the top of its head and light pink “ spectacles ” surrounding the eyes. Unlike blackfurred newborn infants of other baboons, infant Kinda Baboons often have white fur. Unlike Yellow and Chacma baboons, Kinda Baboons usually carry their tails in a simple, gentle curve rather than “ broken. ” Habitat. Light woodland, miombo (Brachystegia, Fabaceae) woodland, and gallery forest.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE3FFE7FA346E24FB0EF786.taxon	food_feeding	Food and Feeding. No systematic studies have been conducted on the ecology and behavior of Kinda Baboons, but they are probably omnivorous with a preference for fruits.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE3FFE7FA346E24FB0EF786.taxon	breeding	Breeding. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE3FFE7FA346E24FB0EF786.taxon	activity	Activity patterns. Kinda Baboons are diurnal and mainly terrestrial.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE3FFE7FA346E24FB0EF786.taxon	biology_ecology	Movements, Home range and Social organization. Preliminary observations suggest that Kinda Baboons, like other baboon species, live in multimale-multifemale groups and do not form spatially coherent one-male units. Dispersal of Kinda Baboons has yet to be determined, but available genetic evidence suggests male dispersal and female philopatry. In Yellow, Chacma and Olive (P. anubis) baboons, females form close social bonds with other females, and most adult grooming partnerships occur between females. To a lesser extent cross-sex “ friendships ” do occur outside of the mating context, and these are identified by grooming partnerships of an adult female and adult male in which the female typically grooms the male. In contrast, adult male Kinda Baboons initiate and maintain proximity to adult females in all reproductive states and, most often, when they have a small infant. Male Kinda Baboons frequently groom females. Male — female relationships and female — female relationships of Kinda Baboons differ substantially from those seen in Yellow, Chacma, and Olive baboons, suggesting that the Kinda Baboon constitutes a further variant of baboon social systems.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE3FFE7FA346E24FB0EF786.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Least Concern on The [UCN Red List (as P. cynocephalus kindae). The Kinda Baboon is listed as vermin in the African Convention on the Conservation of Nature and Natural Resources (as a subspecies of the Yellow Baboon). It possibly occurs in five protected areas: Upemba and Kundelungu national parks in DR Congo and Kafue, Kasanka, and South Luangwa national parks in Zambia. There are no reports of major range-wide threats resulting in a significant population decline of Kinda Baboons.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE2FFE0FFEB6DD2FCF8F7A9.taxon	materials_examined	South Africa, Western Cape Province, Cape of Good Hope.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE2FFE0FFEB6DD2FCF8F7A9.taxon	discussion	In respective contact zones in Zambia, the subspecies griseipes hybridizes with P. kindae and P. c. cynocephalus. The exact distributional boundaries of the subspecies and the extent of hybridization are unclear. The southern and northern populations, carrying genetically different mitochondria (most likely Pw. wrsinus and P. u. gniseipes), geographically overlap in the Loskop Dam Nature Reserve, South Africa, in the east and in central Namibia in the west. Although P. ursinus from sites south of the Limpopo carry “ griseipes ” mitochondria and are quite light in color, they have black hands and feet instead of gray hands and feet as in typical griseipes. A third mitochondrial lineage, most likely representing ruacana, is found in extreme northern Namibia and south-western Angola. Three subspecies recognized.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE2FFE0FFEB6DD2FCF8F7A9.taxon	distribution	Subspecies and Distribution. P. u. wrsinusKerr, 1792 — W & SNamibia, SouthAfrica (exceptNE), Swaziland, andLesotho. However, itisnotclearwhereexactlyitmeetswithgriseipesorruacana. P. u. gniseipesPocock, 1911 — SWZambia, SWZimbabwe, Botswana, SMozambique (SoftheZambeziRiver), andNESouthAfrica. P. u. ruacana Shortridge, 1942 — SW Angola (N along the coast to c ¢. 11 ° 30 ” S) and N Namibia.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE2FFE0FFEB6DD2FCF8F7A9.taxon	description	Descriptive notes. Head-body 68 - 100 cm (males) and 51 - 62 cm (females), tail 42 — 84 cm (males) and 37 - 44 cm (females); weight 25 - 35 kg (males) and 12 - 20 kg (females). Sexual dimorphism in body weight of the Chacma Baboon is pronounced; female body mass is only 52 - 60 % of male body mass. It is the largest of the baboons. Generally, fur is black to dark brown or gray-buff above, with a lighter, almost naked, underside and pale patches on sides of the muzzle. Cape and Kalahari Chacma Baboons are very dark, almost black, but the “ Gray-footed Chacma Baboon ” (P. u. griseipes) 1 s not darker than the Olive Baboon (P. anubis). Adult male Chacma Baboons lack a shoulder mane but have elongated hair tufts along the nape. Facial skin is grayish-black, and both sexes have a set of blackish ischial callosities. Unlike other baboons, the facial part of the skull points downward as well as forward. Nostrils do not protrude beyond the upper lip of the snout. Tail is normally held up and then drops down, “ broken, ” in an inverted “ U. ” In the Southern Cape, the coat of the “ Southern Chacma Baboon ” (P. u. ursinus) is nearly black, along with black hands, feet, and tail. This color becomes lighter in a cline toward the north-east, but extremities are still black. Gray-footed Chacma Baboons are generally fawn colored, with a grayish tail, hands, and feet. The “ Namibian Chacma Baboon ” (P. u. ruacana) is a relatively small subspecies, darker than the Gray-footed Chacma Baboon but tending to be especially dark on the back and crown, where blackened tones may contrast with lighter flanks and limbs. Feet are black. Infants are born with black natal coats that change to adult coloration at 3 - 6 months of age.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE2FFE0FFEB6DD2FCF8F7A9.taxon	biology_ecology	Habitat. A range of habitats including all types of woodland, savanna, steppes, Acacia scrub and semi-desert, swamps, seaside cliffs, and montane regions (up to 3000 m above sea level in the Drakensberg Mountains). As in other baboons, the availability of drinking water limits its overall range in arid areas in Namibia, and, for example. in the Karoo and Kalahari deserts. In some areas, Chacma Baboons benefit from human activities, particularly where water has been provided in cattle troughs and irrigation canals. In some habitats, such as in the Namib Desert, habitat choice is also influenced by predation risk. Cliffs, hills or large trees are necessary night-time retreats. Chacma Baboons also use caves as sleeping sites, in particular during cold southern winters.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE2FFE0FFEB6DD2FCF8F7A9.taxon	food_feeding	Food and Feeding. The Chacma Baboon is an opportunistic omnivore, and its diet varies locally. If available, Chacma Baboons feed on fruits, young leaves, flowers, tuberous roots, bulbs, grasses, bark, twigs, sap, mushrooms, lichens, seeds, shoots, buds, aquatic plants, and seashore animals (e. g. crustaceans, mussels, and snails). Other animal prey is also taken, including invertebrates such as insects and larger vertebrates such as lizards, birds, and small mammals. Occasionally, they may take small antelopes and the young of impala (Aepyceros). Crops (maize, tomatoes, citrus, and root crops) and garbage dumps are raided in settled areas. Lambs and small livestock are taken in some ranching areas. They often beg for food from tourists or take food from houses. Breeding. Births of Chacma Baboons occur throughout the year. The mean length of the menstrual cycle at De Hoop, South Africa, is 39 - 7 days. In Moremi (Okavango), females have their first infant at six years and nine months of age. The interbirth interval is 22 - 39 months, but it is longer in the Drakensberg Mountains. The dominant male of the group tries to monopolize access to sexually receptive females. A male and a female form a temporary consortship, and most mating happens during these consort periods. In contrast to Yellow (P. cynocephalus) and Olive (P. anubis) baboons, a consorting male Chacma Baboon is rarely challenged by other males, and male consort partners do not change often during receptive periods of a female. Males also do not form coalitions to take over a sexually receptive female from the dominant male, as they do in Yellow and Olive baboons. Hence, paternity probability for a particular male Chacma Baboon is higher than in Olive and Yellow baboons, which may be a reason why rates of infanticide are higher in Chacma Baboons than other baboon species. In some populations, more than 30 % of the infants are killed by males. Females give copulation calls; they vocalize during or shortly after copulation. These calls are much louder than those of female Hamadryas (P. hamadryas) or Guinea (PF. papio) baboons.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE2FFE0FFEB6DD2FCF8F7A9.taxon	activity	Activity patterns. Chacma Baboons are diurnal and mainly terrestrial.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE2FFE0FFEB6DD2FCF8F7A9.taxon	biology_ecology	Movements, Home range and Social organization. Depending on ecological conditions and group sizes, the home range sizes of Chacma Baboons vary substantially (1 - 37 km?). Average daily movements are 3 - 6 - 8 - 5 km (range 2 - 13 km). Group size averages 20 - 80 individuals, but it can be as large as 130 individuals. The ratio of males to females is 1: 1 - 5 - 10 - 3. As in Yellow and Olive baboons, the social organization is multimale-multifemale, although Chacma Baboons tend to have smaller groups. Groups are not substructured into one-male units, as they are in Hamadryas Baboons or Geladas (Theropithecus gelada). Solitary one-male units of Chacma Baboons, consisting of a few females and a single male, have been observed rarely, but a survey in the Drakensberg Mountains found that such one-male units made up 50 % of groups above 2100 m. Females remain in their natal groups and form linear dominance hierarchies. Males also establish a dominance hierarchy that usually regulates access to sexually receptive females. Most males emigrate from their natal groups and attain high rank after joining a new group. Male Chacma Baboons may fight vigorously for a dominant position. Before they start fighting, they often have vocalization contests, running up and down producing their specific loud calls, the “ wahoo. ” These contests are believed to signal power and stamina to opponents. As in Yellow Baboons, the social network among related females is the most stable social structure. Grooming and other affiliative behavior occur predominantly within these networks. Studies suggest that these tight relationships provide a fitness advantage for females. Females with a tighter network live longer and reproduce more successfully. Females and males may also form bonds outside sexual consortships, called “ friendships. ” These relationships likely benefit a female via protection for herself and her offspring against sexual harassment and infanticide by other males. The male benefits because it may protect its own offspring or it may invest in future sexual relationship with the respective females. Densities of Chacma Baboons can range from one to more than 24 ind / km? in some protected areas.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE2FFE0FFEB6DD2FCF8F7A9.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Least Concern on The [UCN Red List, including all three subspecies (ruacana under P. ursinus ursinus). The Chacma Baboon is widespread and abundant, and there are evidently no major threats causing a range-wide decline. Outside of protected areas, however, it has been extirpated from large parts of its former distribution, mainly because of agricultural and infrastructural development. Baboons are shot and trapped as crop raiders. Chacma Baboons occur in many protected areas. Southern Chacma Baboons occur in Table Mountain National Park, Namaqua National Park, Mountain Zebra National Park, Augrabies Falls National Park, De Hoop Nature Reserve, Goegab Nature Reserve, Cape nature reserves, Giant’s Castle Game Reserve in South Africa and Namib-Naukluft and Fish River Canyon national parks in Namibia. Gray-footed Chacma Baboons occur in Etosha and Waterberg Plateau national parks in Namibia; Chobe National Park and Moremi Game Reserve in Botswana; Gorongosa National Park and Pomene Game Reserve in Mozambique; Lower Zambezi, Kafue, and Lochinvar national parks in Zambia; and Victoria Falls, Chimanimani, Hwange, and Mana Pools national parks in Zimbabwe. Namibian Chacma Baboons occur in Iona National Park and Namibe Reserve in Angola. It is unclear if baboons in Pilanesberg Game Reserve, Blyde River Canyon Nature Reserve, Hluhluwe-Umfolozi Game Reserve, Ithala Game Reserve, and Kruger National Park (South Africa) are Gray-footed Chacma Baboons.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE5FFE1FFD96D89FC85F8C6.taxon	materials_examined	Africa. Restricted by J. Anderson in 1902 to the Upper Nile.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE5FFE1FFD96D89FC85F8C6.taxon	discussion	P. anubis has regional differences in skull morphology and pelage color. Accordingly, a number of subspecies have been proposed (e. g. tesselatum), but currently none are recognized. Whether some forms would merit subspecies status needs further research. Wherever the distribution of this species encounters that of other baboon species, there are hybrid zones whit strong indications that P. anubis is still in a phase of active expansion. It forms a narrow hybrid zone with P. hamadryas below Awash Falls and elsewhere in northern Ethiopia and central Eritrea. It hybridizes with P. cynocephalus in Tsavo East National Park and Amboseli National Reserve in Kenya. There is a broad clinal hybrid zone of P. anubis x P. cynocephalus between Laikipia district, just to the north-east and east of Mount Kenya, and the Lower Tana River along the Kenya coast. Baboons in this more than 200 km-wide region are intermediate and are difficult to allocate to either P. anubis or P. cynocephalus (baboons become increasingly “ yellow-like ” in their phenotypes toward the Kenya coast). P. anubis from Eritrea and Ethiopia carry maternally inherited mitochondria that are genetically closely related to P. hamadryas mitochondria. P. anubis in Kenya and Tanzania carry mitochondria closely related to P. cynocephalus mitochondria, indicating that male introgression by P. anubis might be the major form of hybridization. P. anubis x P. cynocephalus are also found elsewhere sporadically across the species contact zone in Kenya and Tanzania. It is possible that P. anubis has caused distributions of neighboring baboon species to contract. Hybridization with P. papio is not known for certain but is probable. Even hybridization with Theropithecus gelada has been observed in the wild. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE5FFE1FFD96D89FC85F8C6.taxon	distribution	Distribution. Throughout the N savanna belt from Mali, Guinea, and Sierra Leone to W Eritrea and Ethiopia, and S into N & E DR Congo and NW Tanzania; isolated populations inhabit the Tibesti, Air and Ennedi massifs in the Sahara.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE5FFE1FFD96D89FC85F8C6.taxon	description	Descriptive notes. Head-body 55 - 90 cm (males) and 50 - 70 cm (females), tail 41 - 60 cm (males) and 38 - 46 cm (females); weight 22 - 30 kg (males) and 14 - 18 kg (females). Sexual dimorphism in the body weight of the Olive Baboons is pronounced; female body mass is only 52 - 55 % of male body mass. Although variable in size and coloration, it is generally dark olive-brown above with black hands and feet. Pelage has a “ pepper-and-salt ” or speckled appearance. Adult male Olive Baboons have a mane that is restricted to the foreparts and grades into the shorter body fur (i. e. not sharply set off). Skin of the face and around the buttock pads is purplish-gray to blackish, and the bare area on the rump is smaller than in Hamadryas (FP. hamadryas) or Guinea (P. papio) baboons. Nostrils are pointed and extend a little beyond the snout. Tail is carried directed up for the proximal one-fourth, then falls away in a characteristic “ broken ” fashion, similar to the tail carriage in the Yellow Baboon (PF. cynocephalus). Ischial callosities in males form a single ridge, but in females, they are separated by the genitals. As in other baboons, female Olive Baboons develop prominent periovulatory sexual swellings. Infants are born with black natal coats that change to the adult coloration at 3 - 6 months of age.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE5FFE1FFD96D89FC85F8C6.taxon	biology_ecology	Habitat. Semi-desert, thorn scrub, and open savanna to rocky hills, high-elevation forest, gallery forest, and rainforest at elevations up to 2500 m — the widest variety of habitats of all baboon species. Water is a limiting factor, and groups of Olive Baboons will travel long distances to find it. They readily live near and use cultivated lands.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE5FFE1FFD96D89FC85F8C6.taxon	food_feeding	Food and Feeding. Like other baboons, the Olive Baboon is an omnivorous opportunist. Its diet varies according to region and season and includes mainly fruits and leaves, along with flowers, tuberous roots, grasses, bark, twigs, sap, bulbs, mushrooms, lichens, seeds, shoots, buds, and animal prey (invertebrates, lizards, birds, and small mammals, as well as other primates such as green monkeys, Chlorocebus). Resin, gum, and bark act as a fall-back foods in dry seasons. Swarming locusts and termites provide an occasional glut. Olive Baboons raid crops and garbage dumps.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE5FFE1FFD96D89FC85F8C6.taxon	breeding	Breeding. Female Olive Baboons reach menarche at 4 - 6 years old and have their first infant at 6 - 7 years old. Menstrual cycle length is 34 - 45 days, and the gestation period is 180 - 188 days. Births occur throughout the year, and interbirth intervals are 22 - 27 months. When a female is sexually receptive, males try to monopolize access to her. A male and a female form a temporary consortship (several minutes to days). Most mating happens during these consort periods. In multimale groups, the consorting male is often challenged by other males, and male consort partners may change several times during a female’s receptive cycle. Female Olive Baboons usually mate with several partners. Females give copulation calls, i. e. they vocalize during and shortly after copulation. Males mature at 7 - 10 years old. Most likely, male Olive Baboons take longer to become adults compared with male Hamadryas Baboons, but mature faster than male Yellow Baboons. Olive Baboons may live up to 30 years in the wild.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE5FFE1FFD96D89FC85F8C6.taxon	activity	Activity patterns. Olive Baboons are diurnal and mainly terrestrial.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE5FFE1FFD96D89FC85F8C6.taxon	biology_ecology	Movements, Home range and Social organization. Home range sizes (4 - 44 km? ®) and daily movements of Olive Baboons vary considerably depending on habitat, group size, and season. Home ranges of neighboring groups overlap, and groups may aggregate temporarily at sleeping sites. Daily movements are usually less than 7 km. Besides food, home ranges contain essential localized resources such as water and sleeping sites. Olive Baboons use large trees and rocks as sleeping sites. Like Chacma (FP. ursinus) and Yellow baboons, Olive Baboons live in multimale-multifemale groups of 20 - 120 individuals. Groups in West Africa tend to be smaller (7 - 40 individuals) than those in East Africa (30 - 120 individuals). The ratio of adult males to adult females varies substantially (1: 1 - 9), and there are usually more females than males. Groups are not substructured into one-male units, as in Hamadryas Baboons or Geladas (Theropithecus gelada), although solitary one-male units have been observed, most frequently in West Africa. Most females remain in their natal groups and form linear dominance hierarchies. High-ranking mothers support their daughters during rank conflicts, resulting in a dominance rank of the daughter just below the mother. Rank positions are “ heritable, ” and dominance relationships among matrilines within a group can remain stable over generations. Most males emigrate from their natal groups at 7 - 13 years of age. Secondary emigration into a third group has also been observed. Males also establish a dominance hierarchy that usually regulates access to receptive females. The dominant male of a group may sire most of the offspring during his tenure, but lower ranking males have been observed to form coalitions to challenge higher ranking males and to take over receptive females. The social network among related femalesis the most stable social structure in the group. Grooming and other affiliative behaviors are exchanged predominantly within these networks. Studies suggest that these tight relationships provide a fitness advantage for females. Females with a tighter network live longer and reproduce more successfully. Females and males may also form bonds outside sexual consortships, called “ friendships. ” These relationships likely benefit a female via protection for herself and her offspring against sexual harassment and infanticide by other males. The male benefits because it may protect its own offspring or it may invest in future sexual relationships with the respective females.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE5FFE1FFD96D89FC85F8C6.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Least Concern on The IUCN Red List. The Olive Baboon is listed as vermin in the African Convention on the Conservation of Nature and Natural Resources. It is very widespread and locally abundant, and although persecuted as a crop raider, trapped for biomedical research, and hunted for bushmeat, there are no major threats believed to be resulting in a range-wide population decline. Nevertheless, human-induced habitat loss and degradation impact Olive Baboons. Introduced infectious diseases affect local populations with high mortality rates. Local populations may be threatened. Isolated populations on the Saharan massifs, in particular, merit further research. Further studies also may reveal that certain forms of Olive Baboons are valid subspecies. Olive Baboons occur in many protected areas: e. g. Pendjari and West of the Niger national parks in Benin; West of the Niger National Park in Burkina Faso; Bénoué, Bouba Ndjida, and Waza national parks, Faro Reserve, and Kimbe River and Mbi Crater game reserves in Cameroon; Zakouma National Park in Chad; Garamba, Kahuzi-Biéga, and Virunga national parks and Tayna Gorilla Reserve in the DR Congo; Awash and Bale Mountains national parks, Stephanie Wildlife Sanctuary, and Menagesha-Suba National Forest in Ethiopia; Bui, Digya, and Mole national parks, Kogyae Strict Nature Reserve, Bomfobiri Willdife Sanctuary, and Willi Falls Reserve in Ghana; Marahoué, Comoé, and Mount Péko national parks in Ivory Coast; Lake Nakuru, Marsabit, Meru, Nairobi, Masai Mara, and Samburu national parks in Kenya; West National Park in Niger; Cross River, Gashaka-Gumti, and Kainji Lake national parks and a number of game reserves and forest reserves in Nigeria; Akagera, Nyungwe Forest, and Volcanoes national parks in Rwanda; Dinder National Park in Sudan; Nimule National Park in South-Sudan; Arusha, Gombe Stream, Kilimanjaro, Lake Manyara, Serengeti, and Tarangire national parks in Tanzania; Kéran and Fazao-Malfacassa national parks and Koué Reserve in Togo; and Bwindi Impenetrable, Kibale Forest, Murchison Falls, and Queen Elizabeth national parks and Budongo Forest Reserve in Uganda.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE4FFE1FFFC6292F748F7D7.taxon	materials_examined	Coast of Guinea.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE4FFE1FFFC6292F748F7D7.taxon	discussion	Along its eastern limits, P. papio may hybridize with the larger P. anubis, but reliable information is lacking about the exact positions of these species’ distributional borders and possible interactions. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE4FFE1FFFC6292F748F7D7.taxon	distribution	Distribution. S Mauritania, Senegal, Gambia, Guinea-Bissau, W Mali, W Guinea, and NW Sierra Leone; its occurrence in W Liberia is highly unlikely. Found in extreme Western Africa, the N boundary of which (in Mali and Mauritania) is not completely known.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE4FFE1FFFC6292F748F7D7.taxon	description	Descriptive notes. Head-body 43 - 86 cm (males) and 35 - 69 cm (females), tail 55 - 70 cm (males) and 43 - 56 cm (females); weight 17 - 26 kg (males) and 10 - 14 kg (females). The Guinea Baboon is a medium-sized baboon, with a uniformly reddishbrown body. Face and hands are blackish-red, upper eyelids are white, and ischial callosities are large and pinkish-gray. The Guinea Baboon is strongly sexually dimorphic, with males larger than females. In Niokolo-Koba National Park, Senegal, female body mass is, on average, only 59 % of male body mass. Males also have a well-developed mane or cape on their shoulders that contrasts with shorter fur on the rump and limbs. Tail loops up and back evenly. Nostrils protrude beyond the end of the snout. Females develop sexual swellings of the perineal skin when sexually receptive.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE4FFE1FFFC6292F748F7D7.taxon	biology_ecology	Habitat. Humid coastal mangrove and Guinean forest to arid Sahelian steppe. Guinea Baboons occur in evergreen gallery forest, secondary forest, dry forest, light woodland savanna, and shrubland.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE4FFE1FFFC6292F748F7D7.taxon	food_feeding	Food and Feeding. Similar to other baboons, Guinea Baboons are omnivorous, with fruits as their main food item. They also eat young leaves, seeds, grasses, flowers, tuberous roots, and animal prey (including invertebrates and vertebrates such as lizards, birds, and small mammals, e. g. young bushbuck, Tragelaphus). Guinea Baboons living near farms and plantations will eat rice, maize, yams, groundnuts, and other cultivated crops.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE4FFE1FFFC6292F748F7D7.taxon	breeding	Breeding. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE4FFE1FFFC6292F748F7D7.taxon	activity	Activity patterns. Guinea Baboons are diurnal and mainly terrestrial.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE4FFE1FFFC6292F748F7D7.taxon	biology_ecology	Movements, Home range and Social organization. Information on Guinea Baboons is scarce compared with other baboon species. Most data come from captive groups and from a few short-term field studies in Niokolo-Koba National Park. Their movements vary seasonally between 1000 - 13,000 m / day (mean 4400 m), and home range size is ¢. 3800 ha. Guinea Baboons most likely live in a multilevel or modal social organization, with a complex fission-fusion dynamic on a daily and seasonal basis. The largest aggregations of more than 300 individuals are observed during the rainy season. Observations of captive Guinea Baboons suggest that the basal social unit is the one-male unit (OMU, one adult male and several females and their offspring), similar to but not as strict as that of the Hamadryas Baboon (P. hamadryas). The rigid herding behavior of male Hamadryas Baboons has not been observed in Guinea Baboon: s. In free-ranging Guinea Baboons, the OMU-based organization is not as obvious. In many cases, subgroups or “ parties ” contain more than one adult male, and females interact with several males. Several parties seem to form the next higher level of their society — a “ gang. ” In contrast to other baboons, adult male Guinea Baboons seem to be highly tolerant of each other, they interact affiliatively, and show an elaborate greeting behavior. They have the smallest testes of all the baboons suggesting that sperm competition among males is negligible. Guinea Baboons in Niokola-Koba usually sleep in large trees in gallery forests. In a preliminary population-level genetic study in Niokolo-Koba, overall genetic relationships among group males were similar to those among females, suggesting that males have a similar tendency to stay in their natal group (male philopatry) as do females. In Niokolo-Koba, densities are 2 - 15 ind / km?.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE4FFE1FFFC6292F748F7D7.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Near Threatened on The IUCN Red List. The Guinea Baboon is listed as Class B in the African Convention on the Conservation of Nature and Natural Resources. In the past, large numbers were exported for laboratory use, particularly from Senegal. There is reason to believe that Guinea Baboons have undergone a range contraction because of large-scale agricultural expansion, persecution, and hunting, possibly ¢. 20 - 25 % in the past 30 years. Their adaptability to a wide variety of habitats has probably enabled them to remain locally common, but outside protected areas their numbers have sharply declined because of bushmeat hunting (Mali, Guinea, and Guinea Bissau) or persecution because of their crop raiding (Senegal). Guinea Baboons occur in several national parks: Kiang West in Gambia, Niokolo-Koba in Senegal, Outamba-Kilimi in Sierra Leone, Badiar and Haut Niger in Guinea, and Bafing in Mali. Niokolo-Koba National Park was classified as a World Heritage Site in Danger by the United Nations Educational, Scientific and Cultural Oranganization in 2007.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE4FFE2FAC56D63F6E1F92F.taxon	materials_examined	Egypt.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE4FFE2FAC56D63F6E1F92F.taxon	discussion	P. hamadryas hybridizes with P. anubis in a narrow zone below Awash Falls and elsewhere in northern Ethiopia and central Eritrea. There are no consistent differences between African and Arabian populations. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE4FFE2FAC56D63F6E1F92F.taxon	distribution	Distribution. SW Saudi Arabia (up to Jeddah), W Yemen, NE Sudan (Red Sea Hills), E Eritrea, Djibouti, NE Ethiopia, and N Somalia.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE4FFE2FAC56D63F6E1F92F.taxon	description	Descriptive notes. Head-body 70 - 95 cm (males) and 50 - 65 cm (females), tail 42 — 60 cm (males) and 37 - 41 cm (females); weight 16.4 - 21.3 kg (males, up to 30 kg in captivity) and 10 - 15 kg (females). Male and female Hamadryas Baboons differ markedly in size and physical appearance; female body mass is 58 % of male body mass. Males are ashy orsilvery-gray, with an enormous shoulder cape, mane, or mantle (in older individuals), reaching back to the rump and contrasting with very short hairs on the body and limbs. The mane bushes out to form white cheek whiskers, offset by darker and shorter crown hairs. Adult male Hamadryas Baboons are also typified by their massively built face and dagger-like upper canine teeth. Hands and feet are dark, and tail is tufted with white and held in a simple backward-pointing curve. Skin offace and earsis bright pinkish-red, as is the cushion-like skin around the buttock pads. H. Kummer in 1968 described an east-west distributional gradient in male face color in Ethiopia, with darker faces in the west and more light-red faces in the east. Females and young are drab olive-brown without a mane, and their faces and ears are grayish or dull flesh-colored, resembling females and juveniles of other baboon species. Skin of ischial callosities is similarly colored but not cushion-like. Neonates have light pink faces and black fur. Their fur changes color at 3 - 6 months old. Albinism has been reported (as evidenced by a painting of an albino male by the English artist George Stubbs).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE4FFE2FAC56D63F6E1F92F.taxon	biology_ecology	Habitat. Rocky desert and semi-desert steppe, arid lowland savanna, cliff faces, and alpine grass meadows to 2600 m above sea level. In Eritrea, Hamadryas Baboons also use densely wooded areas (Semenawi Bahri). They appear to be seasonally migratory in at least some parts of their distribution in Ethiopia, where groups may move up into neighboring mountainous areas (up to 3300 m in Simien Mountains National Park) in the wet season. They are dependent on water and never found far from water sources. Hamadryas Baboons are sometimes called “ desert baboons, ” but this name seems to be unjustified because they occur in various habitats from dry coastal lowlands to humid montane forests.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE4FFE2FAC56D63F6E1F92F.taxon	food_feeding	Food and Feeding. Like all baboons, Hamadryas Baboons are opportunistic omnivores. Important foods include grass, grass seeds, rhizomes, roots, tubers, shoots, fruits (notably heglig Balanites, Zygophyllaceae, and buffalo thorn Ziziphus, Rhamnaceae), young leaves, Acacia (Fabaceae) flowers, doum palm fruits (Hyphaene thebaica, Arecaceae), and animal prey (including insects and scorpions and small vertebrates such as lizards and small mammals). During daily foraging, they may move as much as 10 km, and they often make daily climbs of several hundred meters when they descend from their sleeping sites into deep valleys to drink and climb back up again in the afternoon to reach their sleeping cliff. In many places, the foraging behavior and diets of Hamadryas Baboons have changed in recent years because they use crops, garbage dumps, and most importantly, massive stands of the non-native prickly pears (Opuntia sp. Cactaceae). This plant provides a year-round food source, and because of its high water content, it is an additional source of water — a big advantage because access to water is a limiting factor in most parts of their distribution.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE4FFE2FAC56D63F6E1F92F.taxon	breeding	Breeding. Female Hamadryas Baboons reach sexual maturity at 4 - 5 years of age and give birth to theirfirst surviving infant at 5 - 7 years. Males reach adult size at 6 - 9 years of age but are capable of siring infants even earlier, although they have not yet developed the full attributes of an adult male (e. g. silvery gray shoulder mantle). Females have a 30 - 40 day menstrual cycle. They exhibit a pink sexual swelling of the perineal skin when approaching ovulation. Its size fluctuates with concentrations of sexual hormones, reaching its peak around ovulation. Mating and births occur throughout the year, although a weak seasonal birth peak has been found in some studies. In Hamadryas Baboons, it is typically only the adult leader male of a one-male unit that mates with the females. The gestation period lasts 165 - 184 days. Usually only a single young is born. Interbirth intervals average 22 months, but they tend to be much shorter in captivity. In the first weeks after birth, an infant is carried on its mother’s belly, and later it rides “ jockey-style. ” Offspring are nursed for 8 - 12 months. Instances of infanticide have been reported in free-ranging and captive populations. Life span in captivity is typically up to 30 years.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE4FFE2FAC56D63F6E1F92F.taxon	activity	Activity patterns. Hamadryas Baboons are diurnal and terrestrial.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE4FFE2FAC56D63F6E1F92F.taxon	biology_ecology	Movements, Home range and Social organization. Home range sizes are 10 - 40 km? and may overlap extensively with those of other groups. Daily movements can be up to 20 km but are usually 6 - 8 km. Hamadryas Baboons normally spend nights on steep rocky cliffs, but they may also use trees on rare occasions. Their social system is characterized by a multilevel or modular social organization, consisting of three to five levels, superficially resembling the social system of the Gelada (Theropithecus gelada). The smallest social unit is the one-male unit or OMU, comprising an adult “ leader ” male, several females (1 - 9, typically 2 - 3) and their dependent offspring and sometimes one or two “ follower ” males. Follower males socialize with unit members, but mating is usually monopolized by the leader male. Several OMUs and additional “ solitary ” males (males that are not leaders or followers) form a clan. It is believed that clans are male kin-based. Several clans, OMUs, and solitary males together comprise the next higher level oftheir organization, the “ band, ” which can have more than 100 members. Individuals in a band share a common home range, largely synchronize their travel pattern, and use sleeping cliffs together. The band is mostlikely analogous to the multimale-multifemale groups of other baboon species. The largest grouping is the “ troop, ” which is an unstable aggregation of several bandsat sleeping sites, totaling several hundred individuals. At Filoha, Ethiopia, subsets of OMUs within clans suggest a fifth level of society in this population. In contrast to other baboon species where males usually leave their natal groups after reaching maturity, female Hamadryas Baboonsare transferred, often coercively, among OMUs by males within clans and bands, but occasionally also between bands. Males sometimes disperse. Most social behavior occurs within OMUs. Adult males show a ritualized greeting behavior (“ notifying ”), and they cooperate to coordinate their daily travel. The strongest social bonds are between adult females and their leader male within OMUSs, which is in contrast to the strong female — female bonds in other baboon species. Males herd their females aggressively, routinely chasing females and using visual threats and physical aggression (e. g. neck-bites) to control female movement and maintain OMU cohesion.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE4FFE2FAC56D63F6E1F92F.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Least Concern on The [UCN Red List. The Hamadryas Baboon is classified as vermin in the African Convention on the Conservation of Nature and Natural Resources. Prior to 1500, its distribution extended into Egypt. The Hamadryas Baboon is of African origin, but how and when they dispersed into the Arabian Peninsula is still an open question. It is believed that they either crossed the Red Sea via a land bridge near Bab-el-Mandeb during glacial lowsin sea level, or they were transported by ancient Egyptians trading by sea with the Land of Punt. Hamadryas Baboons are rarely hunted, although the Oromo people traditionally use their skins for ceremonial wear. They are chased and occasionally shot or killed with poison as crop pests. In Eritrea, fat of the buttocks of Hamadryas Baboons is used in traditional medicine as a cure for sprain and pulled muscles. They were formerly trapped in large numbers for medical research; a large colony of Hamadryas Baboons, for example, was established 1927 in Sukhumi, Georgia (formerly USSR). The Hamadryas Baboon occurs in several protected areas: Awash National Park, Harar Wildlife Sanctuary, and Simien Mountains National Park in Ethiopia and Asir National Park in Saudi Arabia. It is also found in the proposed Yangudi Rassa National Park, a number of wildlife reserves in the lower Awash Valley in Ethiopia, and several proposed reserves in Eritrea. Part of the population in Awash National Park and other areas along the distributional border of the Hamadryas Baboon in Ethiopia and Eritrea are P. hamadryas x P. anubis (the Olive Baboon) hybrids. As in other baboon species, the conservation status of the Hamadryas Baboon is geographically ambivalent and needs a proper reassessment. Although locally abundant, they are seen as pests by local people and governments, and they have been extirpated from large parts of their former distribution, resulting in a significant population decline. The major threat is loss of habitat due to agricultural expansion and irrigation projects. In a mainly arid environment, many places where water could be found were converted into irrigated agriculture in the last 20 years, and given the dependence of Hamadryas Baboons on water, they were eliminated from these areas.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE7FFE3FA3E62F3F8CCF841.taxon	materials_examined	Ethiopia, Haremat, Semyen, and Gojjam.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE7FFE3FA3E62F3F8CCF841.taxon	discussion	There is a need for further research on the intraspecific taxonomy of 7. gelada, especially the distributional ranges of subspecies. A possible third subspecies was discovered in 1989 in Arsi Province, but it has yet to be formally described. This “ Arsi Gelada ” is possibly synonymous with the described form senex and is found in south-western Ethiopia (Arsi Province) east of the Rift Valley along the Webi-Shebelle Gorges. Two subspecies recognized.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE7FFE3FA3E62F3F8CCF841.taxon	distribution	Subspecies and Distribution. 1. g. gelada Ruppell, 1835 — Ethiopia (N of Lake Tana and W of the Takezé River). 1. g. obscurus Heuglin, 1863 — Ethiopia (S of Lake Tana and E of the Takezé River).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE7FFE3FA3E62F3F8CCF841.taxon	description	Descriptive notes. Head-body 69 - 75 cm (males) and 50 - 65 cm (females), tail 46 — 50 cm (males) and 33 - 41 cm (females); weight 20 - 30 kg (males) and 12 - 16 kg (females). The Gelada is a large, heavily built primate. Both sexes are covered with thick, sooty yellow-brown fur dorsally, with darker limbs and a grayish white underside. Muzzle is rounded and marked by deep lines, and nostrils are unusual in being upturned and set well back on the face. Adult male Geladas are almost twice the size of females (female body mass is 59 % of male body mass) and have a heavy mane around their head and shoulders, which in older individuals hangs almost to the ground. Adult males also have long, curving cheek whiskers and pale eyelids, in stark contrast to the otherwise dusky face. Buttock pads are purplish-gray and rather flat; they do not fuse across the midline in adult females, as they do in adult males. Skin below the buttock pads is bare (edged with vesicles in the female) and incorporates a pair offatty pads below. Tail is shorter than the head-body length and thickly tufted at the tip. Perhaps the most interesting feature in both male and female Geladas, however, is the conspicuous patch of pinkish-red skin on the chest (heart-shaped in males, hourglassshaped in females, and edged with white or gray fur) and an additional, smaller crescent on the throat. Mammae are close to the midline. Hands are ideal for all manner of precise manipulation; they are equipped with highly opposable thumbs and finely proportioned (though notably shortened) index fingers, believed to be an adaptation for manual “ grazing. ” In the “ Northern Gelada ” (T. g. gelada), the mane is dark brown, while the area surrounding the chest patch (which is not extensive) is iron-gray. The “ Southern Gelada ” (1. g. obscurus) is similar to the Northern Gelada, but it has a darker mane and extensive, pure white fur surrounding the chest patch.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE7FFE3FA3E62F3F8CCF841.taxon	biology_ecology	Habitat. High montane regions including grassland, scrubland, and some types of woodland at elevations of 1400 - 4400 m. Steep cliffs provide sleeping roosts. Geladas feed mainly on the flat margins of high grass plateaus, with Agrostis and Festuca grasses (both Poaceae) and giant Lobelia (Campanulaceae) groves. Geladas are poor tree climbers and are almost entirely terrestrial, spending 99 % of their time on the ground. This is partly a consequence of their extreme dietary specialization as a grazer.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE7FFE3FA3E62F3F8CCF841.taxon	food_feeding	Food and Feeding. Grasses are the dietary staple of Geladas, supplemented with roots, bulbs, flowers, seeds, leaves, fruits, and animal prey (including insects). Geladas feed primarily on leaves of grasses. They show some morphological and physiological characteristics that are believed to be adaptations to their grass diet. When feeding, individuals typically shuffle along on their well-cushioned rumps (referred to as “ shuffle-gait ”) plucking blades of grass and occasionally digging around with the help of their long sharp fingernails. During dry seasons when there is heavy overgrazing by livestock, or when Gelada bands are very concentrated in one area, subterranean stems and rhizomes are also excavated. Fruits and invertebrates are eaten opportunistically, and cereal crops may be taken where agriculture encroaches into their habitats.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE7FFE3FA3E62F3F8CCF841.taxon	breeding	Breeding. Female Geladas have a 32 - 36 day reproductive cycle, and the peak is marked by an outline of whitish, fluid-filled beads around the chest and genitals. The gestation period has not been exactly determined, although it appears to be 175 - 188 days, as in other baboons. One, rarely two, offspring are born per pregnancy. Some studies found birth peaks in July-December, but others did not find any seasonality of births. Offspring are nursed for well over a year, during which time the mother seems unable to reproduce again. Young Geladas are uniformly dark smoky-brown. Cases of infantcide have been reported. Expected life span is less than 14 years in the wild, although some individuals may live up to 20 years.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE7FFE3FA3E62F3F8CCF841.taxon	activity	Activity patterns. Geladas are diurnal and terrestrial.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE7FFE3FA3E62F3F8CCF841.taxon	biology_ecology	Movements, Home range and Social organization. Geladaslive in a multilevel or modular society with one-male units (OMUs) as the basal social entity, superficially similar to the social system of Hamadryas Baboons (Papio hamadryas). The OMUs contain one adult “ leader ” male and 1 - 12 (related) females (averaging 3 - 4) with their dependent offspring. Often, 1 - 5 additional males are attached to an OMU, called “ follower ” males. Males that are not leaders or followers of an OMU gather together and form relatively stable all-male units (AMUs). Two to 30 OMUs and an AMU comprise a band, which may have 30 - 260 individuals. Males from AMUs challenge OMU leader males from time to time, trying to take over a unit of females. During takeovers, the risk of infanticide is particularly high. There is even evidence that pregnant females abort if a takeover occurs. If the number of females in an OMU becomes too large, they sometimes split into two or more “ daughter ” units, which retain relatively close social relationships. The OMUs with such close social relationships are called “ teams. ” Bands stay within 2 km of escarpment edges, where they retreat at night or if alarmed. As a result, home ranges are linear, encompassing as little as 1 - 3 km? for a band’s core area (although their annual home range can be 70 km? ®). In general, Gelada bands are less stable in their composition than are those of Hamadryas Baboons; they are best regarded as a set of OMUSssharing a foraging area. At times, several bands may aggregate at sites where food is abundant. Up to 1000 Geladas can then be seen grazing together. Surveys in a number of areas suggest overall densities of 15 - 60 ind / km?, but densities within home ranges commonly exceed 70 ind / km? ®.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE7FFE3FA3E62F3F8CCF841.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Least Concern on The [UCN Red List, including the subspecies obscurus, but the nominate subspecies gelada is classified as Vulnerable. The Gelada is listed as Class A in the African Convention on the Conservation of Nature and Natural Resources. Male Geladas have been regularly hunted by local Ethiopians, who use their enormous manes to make ceremonial headdresses. In the past, the numbers of adults may have been reduced as a result of selective shooting for their capes. There are historic records of capes being made into fur hats for tourists, but that no longer occurs because it is now extremely difficult for any tourist to leave the country with items made from Gelada skins. Although hunting may no longer be the main threat, they are still chased and shot as crop pests. Geladas have been trapped as laboratory animals in the past (e. g. 1200 individuals were imported into the USA in 1968 - 1973). Currently, the main threatis deterioration of their habitat and competition with domestic livestock and agriculture, resulting in significant habitat loss. Northern Geladas are known to occur in one protected area, Simien Mountains National Park, but Southern Geladas do not occur in any protected area. The proposed Blue Nile Gorges National Park and Indeltu (= Shebelle) Gorges Reserve would serve to protect a large part of the total population, which is currently estimated to be less than 250,000 individuals (more than 440,000 in the 1970 s). The Indeltu Gorges Reserve would protect the unnamed subspecies.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE8FFEEFFF46DDEFE2AFE4A.taxon	materials_examined	Upper Congo.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE8FFEEFFF46DDEFE2AFE4A.taxon	discussion	Molar flare and ischial callosities that fuse across the midline are two physical conditions unique to the genus Allenopithecus among Cercopithecini, but they are shared with some, if not all, Papionini. Another feature common to most Papionini is the periodic sexual swelling of females, shared only with Miopithecus among the Cercopithecini. Recent molecular studies suggest a tribal affiliation of Allenopithecus with the Cercopithecini rather than the Papionini. It is the sister taxon ofall the Cercopithecini. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE8FFEEFFF46DDEFE2AFE4A.taxon	distribution	Distribution. NW DR Congo and NE Republic of the Congo, in lowland forests of the Congo Basin on both sides of the lower and middle Congo (c. 16 ” E to 26 ° - 27 ° E); S of the Congo Riverits distribution extends E to the Lomami River system (c. 3 ° N to 6 ° 30 ° S); it has also been recorded on twoislands in the Sangha River. Its presence in NE Angola requires confirmation.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE8FFEEFFF46DDEFE2AFE4A.taxon	description	Descriptive notes. Head-body 45 - 50 cm (males) and 40 - 45 cm (females), tail 51 cm (males) and 36 cm (females); weight 5.9 - 6.1 kg (males) and 3.2 - 3.7 kg (females). Male Allen’s Swamp Monkeys are larger than females. They are very robust, with short limbs, a relatively short tail, and broad hands and feet. They have pointed macaquelike ears and a dark face, with pale, off-white eyelids and chin. Facial hair is agouti black-yellow, giving a dark olive — green effect; it is especially long on cheeks and forms a circular face mask bordered with narrow black eyebrow bands extending back to ears. Crown appears like a broad, flattened cap. Underside is grayish, off-white, slightly speckled; throat is gray but not speckled. Male perineum has a red margin and an orange perineal tuft; scrotum is bluish white. Males have a scent gland on their chest that is marked by bristly white hair. Digits on feet are long and powerful, and hands are shorter and broader than is typical of guenons. Neonatal coatis yellow-brown, with an almost white crown.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE8FFEEFFF46DDEFE2AFE4A.taxon	biology_ecology	Habitat. Swamps in lowland moist forests, bordering lakes and rivers that are seasonally flooded. Allen’s Swamp Monkey is largely arboreal when the forest is flooded but more terrestrial when flooding recedes. It is also found in mangroves and occasionally enters terra firma forests and more open formations. It enters fields to raid crops.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE8FFEEFFF46DDEFE2AFE4A.taxon	food_feeding	Food and Feeding. A study of Allen’s Swamp Monkeys in 1985 reported a diet of 81 % fruit, 17 % invertebrate prey, and 2 % miscellaneous items such as flowers, nectar, pith, and roots. They lick nectar of, and thereby pollinate, Daniellia pynaertii (Fabaceae) flowers. They also eat crabs and scoop fish out of muddy pools during the dry season.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE8FFEEFFF46DDEFE2AFE4A.taxon	breeding	Breeding. Births of Allen’s Swamp Monkeys have been recorded throughout the year. Life span is estimated to be 28 years.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE8FFEEFFF46DDEFE2AFE4A.taxon	activity	Activity patterns. Allen’s Swamp Monkeys are diurnal and semi-arboreal-semi-terrestrial. Locomotion is quadrupedal, with climbing and leaping. They use the lowerstrata of the forest canopy and often forage on the ground. They can swim and will dive into a river to avoid predators, or flee on the ground rather than climb to safety. They sleep in trees 8 - 20 m above the ground along river edges and lake borders.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE8FFEEFFF46DDEFE2AFE4A.taxon	biology_ecology	Movements, Home range and Social organization. Home range sizes and the lengths of the daily movements of Allen’s Swamp Monkeys are unknown. They live in multimale-multifemale social groups of 23 - 57 individuals, but it is possible that the social organization is one of harems. Groups of 2 - 8 individuals tend to travel and forage together. Single males may travel separately during the day but join up with the group at night. They produce baboon-like grunts as they travel and also guenon-like, highpitched alarm chirps.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFE8FFEEFFF46DDEFE2AFE4A.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Least Concern on The [UCN Red List. Allen’s Swamp Monkeyis listed as Class B in the African Convention on the Conservation of Nature and Natural Resources. They are hunted from boats on rivers at night as they sleep in trees overhanging the river, but hunting is not believed to be so serious that it results in a decline that would warrant listing it in a threatened category. Further study is required, however, to assess the effects of hunting. Allen’s Swamp Monkeys are found in relatively high densities in Salonga National Park and Tumba-Lediima Reserve, DR Congo, and are marginally present in Nouabalé-Ndoki National Park in the Republic of the Congo. It is a little-known species that is very difficult to observe and follow in the wild, and further information on its distribution and conservation status is needed.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEBFFEEFA2B64A1F6CEF277.taxon	materials_examined	No specific type locality given. Restricted by C. P. Groves in 2001 as “ endemic to the equatorial coastal watersheds between Cabinda and the River Nyong. ”	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEBFFEEFA2B64A1F6CEF277.taxon	discussion	The genus Miopithecus was considered monotypic, ranging from southern Cameroon to northern Angola, until 1969 when A. B. Machado showed that there were two species: one north and the other south of the Congo River. J. Kingdon described M. ogouensis as the form to the north of the Congo River, with the Ogooué River Basin at the center of its distribution. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEBFFEEFA2B64A1F6CEF277.taxon	distribution	Distribution. Equatorial W Africa from S Cameroon (S of the Sanaga River) to NW Angola (Cabinda), including Equatorial Guinea, Gabon, and the W Republic of the Congo; there is what would appear to be an outlying population along a N tributary of the Sanaga, the Djerem River and one ofits tributaries, at ¢. 6 ° N.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEBFFEEFA2B64A1F6CEF277.taxon	description	Descriptive notes. Head-body 34 cm (males) and 28 cm (females), tail 43 cm (males) and 37 cm (females); weight 1 - 4 kg (males) and 1 - 1 kg (females). The Northern Talapoin Monkey is the smallest monkey in Africa. Its head is relatively large compared to its overall size, thus giving it a juvenile appearance. Dorsal surfaces are yellowish olive-green, speckled with gray, and slightly tinged with yellow on limbs. Underside is creamy white, and tail is grayish khaki, darker toward the tip. Face is covered by golden-yellow whiskers. Muzzle, skin around eyes, and ears are pinkish, and nose is gray. Fingers and toes are grayish-pink. Scrotum is large and pale blue. Females have cyclical pink perineal swellings during the periovulatory period.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEBFFEEFA2B64A1F6CEF277.taxon	biology_ecology	Habitat. Never more than a few hundred meters away from rivers and lakes in lowland equatorial riparian forest, swamps, seasonally flooded forest, or periodically flooded mangroves and coastal forests. Dense vegetation helps hide these small monkeys from predators. Talapoins are able to swim, and they will jump into a river and swim underwater to escape from predators.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEBFFEEFA2B64A1F6CEF277.taxon	food_feeding	Food and Feeding. Northern Talapoin Monkeys eat fruits, seeds, and insects, in addition to agricultural crops, bird eggs, and even honey. Fruit makes up ¢. 80 % of the diet, and favored fruits include plums, figs (Ficus, Moraceae), umbrella trees (Musanga cecropioides, Urticaceae), African grape (Pseudospondias microcarpa, Anacardiaceae), African ginger (Aframomum, Zingiberaceae), and flesh of oil palm nuts. They eat beetles, grasshoppers, caterpillars, butterflies, and spiders; bird eggs are taken opportunistically. The Northern Talapoin Monkey is also known to attack nests of some bees to eat honey or the brood. By preying on nests built above rivers, they are able to plunge into the water to avoid attacks by the disturbed insects. The Northern Talapoin Monkeyis also attracted out of the forest to feed on agricultural crops including, bananas, pawpaw, corn, maize, cucurbits, peanuts, and cassava. Leaves are rarely eaten.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEBFFEEFA2B64A1F6CEF277.taxon	breeding	Breeding. Reproduction in the Northern Talapoin Monkey is very seasonal. Females give birth during the short dry season (November — April) after a 5 - 5 month gestation. It appears that some females are able to give birth annually, and in large troops more than 20 infants may be born within two months. Females reach sexual maturity at ¢. 3 — 4 years of age and exhibit a genital swelling when sexually receptive.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEBFFEEFA2B64A1F6CEF277.taxon	activity	Activity patterns. Both species of talapoins are very agile when moving through dense undergrowth; they are able to jump and leap with great precision. When climbing along thin vines, the tail is used as a counterbalance. When moving along the ground, they walk almost tripedally, using the tail as a third foot. They are also confident swimmers. Feeding is concentrated into one early morning bout, with another in late afternoon. Each group has a regular sleeping site that it occupies almost every night. Sleeping sites are always beside a river that is both wide and deep enough to allow individuals to escape from predators by dropping orjumping into the water.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEBFFEEFA2B64A1F6CEF277.taxon	biology_ecology	Movements, Home range and Social organization. Northern Talapoin Monkeys form large groups averaging more than 60 and sometimes exceeding 100 individuals. Groups have multiple adult males, with twice as many adult females and their young. Solitary males are very rare. Home ranges are 100 - 500 ha, indicating densities of 40 - 90 ind / km? *. Northern Talapoin Monkeys are very active and travel an average of 2300 m / day in search of food.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEBFFEEFA2B64A1F6CEF277.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Least Concern on The [UCN Red List. The Northern Talapoin Monkeyis listed as Class B in the African Convention on the Conservation of Nature and Natural Resources. It has no major threats. The Northern Talapoin Monkey is small and its choice of habitat deters hunters, who seldom choose to venture into swampy forest to chase after such small game. Their habitat also generally escapes conversion by logging and farming. Human settlements act in their favor because they exploit cultivated fruits ofriverside gardens. Densities of Northern Talapoin Monkeys double close to human settlements, and their overall numbers have probably increased, at least in the past, because of humans. Talapoins are found in a number of protected areas across their distribution.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEBFFEEFF2367D2F75AFE94.taxon	materials_examined	the type locality is unknown.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEBFFEEFF2367D2F75AFE94.taxon	discussion	The genus Miopithecus was considered to be monotypic, with a distribution from southern Cameroon to northern Angola, until 1969 when A. B. Machado showed that it comprised two species: one north and the other south of the Congo River. The typical form, M. talapoin, and all names given to putative subspecies, referred to those individuals south of the river in Angola. |. Kingdon addressed the omission and gave the name M. ogouensis to the talapoin monkeys north of the Congo River. Miopithecus differs from Chlorocebus, Erythrocebus, and Cercopithecus in the distal reduction ofits third upper molar, the high degree of development of the facial gestures, and, as in Allenopithecus, the cyclical sexual swellings of the females. A. Tosi and colleagues in 2004 suggested that Miopithecus is an early evolutionary branch that differentiated after the separation of all the other guenons from Allenopithecus. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEBFFEEFF2367D2F75AFE94.taxon	distribution	Distribution. SW DR Congo (both sides of the Kasai River and along the Mebrige, Loge, Cuanza, Nhia, Cuvo, and Cuangorivers) and NW Angola (from the coastal watersheds of the Congo River S to c. 13 ° S).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEBFFEEFF2367D2F75AFE94.taxon	description	Descriptive notes. Head-body 26 - 45 cm (males), tail ¢. 53 cm; weight 1.2 - 1.3 kg (males) and 0 - 75.0 - 8 kg (females). The Southern Talapoin Monkey is the smallest of the Old World monkeys. Pelage is relatively long and coarsely banded yellow and black. Underside is contrasting white or off-white, and outer surfaces of limbs are yellowish, with less grizzling than on the back. Tail is relatively long; brown above and yellowish gray below; cheek hairs, covering suborbital fossae, are directed downward and backward. There are well-developed preauricular tufts, pointing upward and backward, that are olive yellow with a lighter central part. Ears are large, nose and bordering facial skin are black, giving the appearance of a mask, but the rest of the facial skin is pinkish ocher. Upperlip is lighter in the middle than at the sides. Scrotum is salmon-colored medially, pale blue laterally. Females have pink sexual swellings during their periovulatory periods.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEBFFEEFF2367D2F75AFE94.taxon	biology_ecology	Habitat. Strictly riparian areas: dense evergreen vegetation of gallery forests, seasonally flooded forest, mangroves, and swamps on riverbanks, including those flowing through miombo (Brachystegia, Fabaceae) woodland. Southern Talapoin Monkeys are increasingly found in and near cultivated areas where they benefit from dense secondary forest and reduced presence of predators. They rarely venture more than 450 m from a river. They occur at elevations from sea level to ¢. 700 m.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEBFFEEFF2367D2F75AFE94.taxon	food_feeding	Food and Feeding. Southern Talapoin Monkeys feed mainly on fruits (43 % of the diet in one study) and also seeds, young foliage, flowers, and small animals, including reptiles and invertebrates (arthropods and freshwater shrimp). They also eat bird eggs. They raid crops and have been reported to take manioc roots that are left by farmers to soak.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEBFFEEFF2367D2F75AFE94.taxon	breeding	Breeding. Female Southern Talapoin Monkeys reach sexual maturity at c. 4 years old, and males as late as nine years old. The menstrual cycle is ¢. 35 days. Females display a cyclical sexual swelling. The gestation period is 5 - 6 months. Births are generally of singletons. Average life span is c. 27 years.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEBFFEEFF2367D2F75AFE94.taxon	activity	Activity patterns. Southern Talapoin Monkeys are diurnal and semi-terrestrial-semiarboreal. Locomotion is quadrupedal, with some climbing and leaping. They generally move and forage in lowerlevels of the forest understory and sometimes forage on the ground.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEBFFEEFF2367D2F75AFE94.taxon	biology_ecology	Movements, Home range and Social organization. Groups of Southern Talapoin Monkeys travel 1500 - 3000 m / day. Home ranges are 122 - 500 ha. They live in large multimale-multifemale groups of 40 - 123 members. Groups living in close proximity to humans are reported to be double or even triple the sizes of groups living in more undisturbed forest. They form single sex groups, with males interacting with males, and females with females.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEBFFEEFF2367D2F75AFE94.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Least Concern on The [UCN Red List. The Southern Talapoin Monkeyis listed as Class B in the African Convention on the Conservation of Nature and Natural Resources. There is very little known about its population status, but there are no obvious known threats in its range, and therefore no reason to expect that it would be declining at a rate that would warrant listing in a threatened category. It is generally not hunted because it is so small. Its occurrence in protected areas has not been documented.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEDFFE9FF2B64A0FB7EFA96.taxon	materials_examined	Senegal.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEDFFE9FF2B64A0FB7EFA96.taxon	discussion	Although considered monotypic by most experts, several distinct subspecies have been described, distinguished mainly by slight differences in facial coloration and pattern. Some this variation, has been shown to be related to female reproductive physiology. In late pregnancy, the nose becomes white, black temporal lines disappear, and the face becomes light gray. These changes persist until c. 45 days postpartum. Nevertheless, subspecieslikely do exist, as described below. At least one incidence of hybridization with a Chlorocebus aethiops pygerythrus has been reported in captivity. A fourth subspecies (villiersi) named by Dekeyser in 1950 is recognized by some. It occurs in Niger from Air and north of the bend of the Niger River. Clear demarcation of distributions of these possible subspecies is still unclear. Additional studies are needed. Three subspecies recognized.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEDFFE9FF2B64A0FB7EFA96.taxon	distribution	Subspecies and Distribution. E. p. patasSchreber, 1774 — sub-SaharanAfricafromSMauritaniaandSenegaltoChadandCentralAfricanRepublic; isolatedpopulationsinhabittherockyAirMassifofNNigerandtheEnnediMassif, Chad. E. p. baumstarkiMatschie, 1905 — NCTanzania (SerengetiandAthiPlainsandtheWsideofMt. Kilimanjaro), butthedistributionisnotwelldocumented. E. p. pyrrhonotus Hemprich & Erhenburg, 1829 — Sudan (N to Nubia), South Sudan, W Ethiopia, N DR Congo, N Uganda, and Kenya. Introduced into south-western Puerto Rico in the Caribbean Islands.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEDFFE9FF2B64A0FB7EFA96.taxon	description	Descriptive notes. Head-body 60 - 87.5 cm (males) and 48 - 52 cm (females), tail 63 — 72 cm (males) and 48 - 55 cm (females); weight 7 - 13 kg (males) and 4 - 7 kg (females). The Patas Monkey has substantial sexual dichromatism and dimorphism. Both sexes are reddish-brown above, with underside, rump, neck, chin, inner surfaces of limbs, and outer surfaces of lower limbs white in males, and fawn in females and juveniles. In infancy, crown hairs are mostly black. These begin to change at c. 5 - 6 weeks of age. By 7 - 8 weeks, crown has the normal adult coloring. Adult males have a shaggier, more brightly contrasting coat, with russet tail, hindquarters, and crown, and they often develop a grayish shoulder mane with age. Females and juveniles are often the color of dry grass, with shades of fawn, russet, and gray, making them difficult to see in their savanna habitats. In adulthood, males can be twice the size of females, with some exceptional individuals weighing up to 20 kg. The nominate “ Western Patas Monkey ” (E. p. patas) has a pink face and a dark nasal spot, but more generally, its face ranges from pink through dirty white to black with a variable pattern, short cheek fringes, a white moustache, and a black band across the forehead extending to ears. Penis is red, and scrotum is blue. Ischial callosities are yellowish-white or grayish-white, and separated in both sexes. The status of subspeciesis still debated. Of note, the “ Southern Patas Monkey ” (E. p. baumstarki) has been described as a “ pallid Serengeti isolate, ” while the “ Eastern Patas Monkey ” (E. p. pyrrhonotus) is said to have a blackish face and white nose. Some recognize another subspecies, villiersi, which differs from the Western Patas Monkey only in its smaller size. Further study with a more detailed description and diagnosis is needed to determine whether this is indeed a fourth subspecies.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEDFFE9FF2B64A0FB7EFA96.taxon	biology_ecology	Habitat. Dry, open Acacia woodland savanna, semi-desert, and grassland. The Patas Monkey is mainly terrestrial and may also be found in volcanic rocky outcrops and woodland-grassland margins. They can live at elevations of up to 1900 m, although they are usually found in lower elevations.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEDFFE9FF2B64A0FB7EFA96.taxon	food_feeding	Food and Feeding. Patas Monkeys are omnivores, eating a wide variety offruits, grass seeds, bulbs, leaves, new shoots, berries, flowers, tamarind beans, fungi, and bird eggs. They use many Acacia (Fabaceae) tree parts (fruits, galls, leaves, and gums). Some populations have 20 % gum in their diets. Animal prey is also an important part of the diet of the Patas Monkey. This can include many arthropods, lizards, and nestling birds. Grassitself is rarely eaten. They eat prickly pear cactus (Opuntia, Cactaceae) that has been introduced to Kenya. Patas Monkeys need to drink daily, and they visit rivers, rock pools, waterholes, or puddles in the dry season. Groups occasionally raid cultivated areas. Varying availabilities of cultivated crops and insects result in great seasonal changes in their diet. In general, Patas Monkeys avoid large, tough foods, preferring instead small fruits or berries. The infrequent need for cutting and food preparation corresponds to their relatively small incisors.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEDFFE9FF2B64A0FB7EFA96.taxon	breeding	Breeding. Female Patas Monkeys have a 30 day reproductive cycle. They show no evidence of a sexual swelling during the periovulatory period, which lasts for ¢. 3 - 6 days (range 1 - 10 days). The interbirth interval is c. 1 year, which is shorter than would be expected for a monkey of its body mass. The 2 - 5 month conception period tends to be seasonal and synchronized within the group. Typically, conception occurs in the wet season (summer), and births occur in the dry season (winter). In northern Cameroon, most mating takes place in the wet season and resulting births occur in late December to mid-February in the dry season. Timing varies geographically and is subjectto changes in the environment. One measure of female proceptivity is the unmistakable behavior ofsoliciting copulation by crouching in front of a male and exhaling into her cheek pouches. Despite this display, the female’s behavior does not usually appear to influence copulation rates of individual males. As seen in many other primate species, copulations tend to occur in early-to-late morning; copulations at mid-day and beyond are uncommon. During late pregnancy, the normally black nose of the female turns white, and the remainderof the face becomeslight gray, before ultimately reverting to its natural coloration c. 45 days post-partum. One dark brown or blackish offspring is born after a gestation of 167 days. Births are typically diurnal, unlike most other monkeys. This is considered part of the Patas Monkey’s anti-predator strategy. As a result, several births have been witnessed and fully described by researchers. The entire labor and delivery process is fairly short (less than one hour), and the mothertypically places her newborn on the grass while eating the placenta. Juveniles are light brown with a pink face and are carried ventrally. The juvenile phase is short, with sexual maturity reached at three years or even earlier for females and 4 - 5 years for males. Individuals may live c. 30 years in captivity.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEDFFE9FF2B64A0FB7EFA96.taxon	activity	Activity patterns. Patas Monkeys are diurnal and mainly terrestrial. Although they can easily climb trees when alarmed, they usually rely on their speed on the ground to escape from danger. Their relatively robust forelimb bones are well adapted for living a terrestrial lifestyle. They spend ¢. 60 % of their time on the ground, and a full 90 % of their feeding time is on the ground. They use digitigrade hand and foot postures while walking, which helps increase stride length. Consequently, the Patas Monkey is generally considered to be the fastest primate. Nonetheless, they cannot outrun some of their predators (e. g. Cheetahs, Acinonyxjubatus), so they have a finely tuned system of vigilance and cryptic coloring to aid in predator avoidance. When traveling on the ground, Patas Monkeys tend to remain relatively silent. Daily movement may vary from 500 m to 12,000 m, and the group often splits before sunset. They rarely begin moving before 07: 00 h and usually stop moving for the day by 19: 00 h.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEDFFE9FF2B64A0FB7EFA96.taxon	biology_ecology	Movements, Home range and Social organization. Average group size of the Patas Monkey is 15, although most mixed-sex groups have 20 - 51 individuals. The home range is large (e. g. 51 - 8 km? for one group of 31 individuals), but all-male groups can have twice the home range size of mixed-sex groups. During the night, a group can spread out over as much as 25 ha. Such large home ranges and expansive sleeping patterns serve a protective role against predators but also aid in better exploitation of the habitat. Female Patas Monkeys remain in their natal group throughout their lives, while males disperse at maturity. They may then live alone, as extragroup males, or become the sole resident male of a female group. Their social system is not representative of their mating system; one-male groups are common, but this does not represent a harem unit. During the mating season, other adult males that normally live alone or in “ bachelor ” groupsjoin female groups. For about one month, mating occurs in a rather promiscuous manner; the original resident male does not have an advantage. Some mate guarding is occasionally observed, and there is often fighting among males associated with the mating season. Despite these fights, a definitive hierarchy among males is rarely identified. Adult males engage in jumping and calling as common alarm behaviors to distract predators, thus allowing females and young to escape danger. The role of adult males in an infant's life seems to vary according to study site. Some researchers report that males are often scarce in the group, resulting in an infant being several months old before it is made aware of their presence. Others report keen watchfulness on the part of adult males. In all cases, however, allomaternal behavior is regularly seen. In fact, underfield conditions, it can be difficult to identify which infants belong to which adult females.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEDFFE9FF2B64A0FB7EFA96.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Least Concern on The IUCN Red List. The subspecies have not been assessed. The Patas Monkeyis listed as Class B in the African Convention on the Conservation of Nature and Natural Resources. It remains widespread and relatively common in parts of its distribution, although it is normally found at low population densities. Although their optimum density is estimated to be c. 1 - 5 ind / km?, one study reported as few as 110 Patas Monkeys in 311 km? In Kenya, the current distribution is only 54 % of the known historic (pre- 1995) distribution. Drought and conversion ofsuitable habitat into farmland are growing threats. In fact, there has been a marked decrease in numbers of Patas Monkeys in parts of their range, because of habitat loss from increasing desertification. This results from unsustainable land-use practices, such as overgrazing by cattle and clearing of savanna land for crops. The resulting proximity to farmland leads to predictable crop-raiding and retaliatory persecution by farmers. Patas Monkeys are occasionally hunted by humans for meat (which was long believed by locals to cure leprosy). This has apparently led to a further decline in their numbers; Nigerian hunters report that Patas Monkeys were much more common 15 - 20 years ago. At the species level, the Patas Monkeyis known to occur in 37 protected areas: Pendjari and West of the Niger national parks in Benin; West of the Niger National Park in Burkina Faso; Bénoué, Bouba-Ndjida, Kalamaloué, and Waza national parks in Cameroon; Zakouma National Park in Chad; Abuko Nature Reserve and Kiang West National Park in Gambia; Bui, Digya, Mole national parks and Kogyae Strict Nature Reserve in Ghana; Comoé National Park in Ivory Coast; West of the Niger National Park in Niger; Cross River and Kainji Lake national parks, Bagale Hills, Igangan Ohosu, and Oban Hills forest reserves, Borgu, Kwiambana, Mutum Biya, and Yankari game reserves, and Old Oyo and Upper Ogun reserves in Nigeria; Basse Casamance, Niokolo-Koba, and Djoudj national parks in Senegal; Dinder National Park in South Sudan; Serengeti National Park and Grumeti and Tkorongo game reserves in Tanzania; Keran, Koué, and Malfacassa reserves in Togo; and Kidepo Valley and Murchison Falls national parks in Uganda. Many Patas Monkeys also occur outside protected areas. The Laikipia District in Kenya is said to have the largest population, and they are routinely found in many unprotected areas of Nigeria and Sierra Leone, where bushmeat hunting remains a threat. Therefore, while the Patas Monkey is listed as Least Concern, they should be monitored in areas that are not protected; growing human populations and threatening activity may result in local risks of population decline. Of particular concern is that the Southern Patas Monkey of Tanzania declined by ¢. 33 % from 1995 through 2009. Their area of occurrence has decreased from 30,500 km? to 19,000 km? and population numbers are thoughtto be less than 900 individuals, perhaps even less than 150 individuals. Urgent conservation action is required for this subspecies.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFECFFE9FFEE6368F948FCBC.taxon	materials_examined	Sudan, Sennaar.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFECFFE9FFEE6368F948FCBC.taxon	discussion	C. aethiops intergrades with C. pygerythrus in south-western Ethiopia; it is unclear if the resulting hybrids are the form originally described as C. a. zavattari. Some authors recognize the form matschiei from the western highlands of Ethiopia as a distinct subspecies. It is said to have a more fawn-to-russet back than typical C. aethiops, with the internal aspects of the limbs more gray. Its tail is also shorter, with a reduced terminal tuft and reduced tufts at the base. A more likely possibility is that it is merely a population of C. aethiops that has hybridized extensively with C. djamdjamensis. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFECFFE9FFEE6368F948FCBC.taxon	distribution	Distribution. NE Africa E of the White Nile, in Eritrea, Ethiopia (S to the Omo River and E as far as the Rift Valley), Djibouti, SE Sudan (from Khartoum in the N to Mongalla in the S), and E South Sudan; possibly in S Egypt.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFECFFE9FFEE6368F948FCBC.taxon	description	Descriptive notes. Head-body 42 - 60 cm (males) and 30 - 50 cm (females), tail 46 - 76 cm (males) and 41 - 66 cm (females); weight 3.1 - 6.4 kg (males) and 1.5 - 4.9 kg (females). The Grivet Monkey is grizzled-olive above with a white underside, grayish limbs, and a yellowish-grizzled-olive crown. Face is black, with a fine white moustache and fluffy, elongated cheek fringes, the latter continuous with a narrow white brow band. Hands and feet are pale, with light brown digits. There is a white tuft at the base of the tail. Scrotum is sky-blue.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFECFFE9FFEE6368F948FCBC.taxon	biology_ecology	Habitat. Savanna and open woodland and often forest-grassland mosaic habitats. Grivet Monkeys are normally in close proximity to rivers. They are extremely adaptable and are found in rural and urban environments; the latter may increase risk of problems arising from human-wildlife conflict in cultivated areas.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFECFFE9FFEE6368F948FCBC.taxon	food_feeding	Food and Feeding. Grivet Monkeys eat mainly grasses, fruits, flowers, bulbs, roots, young leaves, eggs, and animal prey. The latter category includes invertebrates, chameleons, fledgling birds, and small mammals such as rodents and even galagos (Galagidae). They are especially fond offigs and other fruiting trees, and in some areas, they are heavily dependent on parts of thorn trees (Acacia, Fabaceae), including seeds, gums, thorns, and bark. Grivet Monkeys raid crops and are considered a major pest in some parts of their distribution. If not hunted, they become habituated to humans and even take handouts from tourists.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFECFFE9FFEE6368F948FCBC.taxon	breeding	Breeding. Female Grivet Monkeys show no sexual swelling. A single offspring is born after a 6 month gestation. Individuals may live more than 30 years.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFECFFE9FFEE6368F948FCBC.taxon	activity	Activity patterns. Grivet Monkeys are diurnal and mainly terrestrial.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFECFFE9FFEE6368F948FCBC.taxon	biology_ecology	Movements, Home range and Social organization. Grivet Monkeys live in groups of 6 - 20 individuals, averaging a dozen. Behavioral and genetic studies indicate that males transfer among groups, while females tend to remain in their natal groups. Like other members of this genus, Grivet Monkeys have a complex social structure; most groups have multiple adult males that are usually outnumbered by adult females.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFECFFE9FFEE6368F948FCBC.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Least Concern on The [UCN Red List. Grivet Monkeys are listed as Class B in the African Convention on the Conservation of Nature and Natural Resources. They are often common and are able to adapt to a wide variety of habitats throughout their distribution. There are presumed to be no major threats. It is known to occur in one protected area: Awash National Park in Ethiopia. Genetic analyses of some populations in Ethiopia indicate a low level of variability compared with many other cercopithecines, suggesting the possibility that populations of Grivet Monkeys have undergone fluctuations in size in recent times as a result of climate change.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFECFFEAFAF46697FB04FBD3.taxon	materials_examined	no type locality.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFECFFEAFAF46697FB04FBD3.taxon	discussion	C. tantalus hybridizes with C. pygerythrus in the DR Congo and Uganda. There also may be some limited interbreeding with C. sabaeus in Ghana. Three subspecies are recognized. Subspecies and Distribution. C. t. tantalusOgilby, 1841 — sub-SaharanAfricafromEBurkinaFasoandGhana (EofVoltaRiver) toCentralAfricanRepublicandNWDRCongo. C. t. budgettiPocock, 1907 — NEDRCongo, Uganda, andNWKenya. C. t. marrensis Thomas & Hinton, 1923 — W Sudan (W of White Nile) and South Sudan from the Imatong Mts in the S to Jebel Marra in the N (where it is isolated from other populations by desert).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFECFFEAFAF46697FB04FBD3.taxon	description	Descriptive notes. Head-body 44 - 83 cm (males) and 30 - 50 cm (females), tail 58 — 60 cm (males) and 41 - 66 cm (females); weight 4.3 - 6.4 kg (males) and 3.4 - 9 kg (females). The Tantalus Monkeyis the largest species of Chlorocebus. Back and crown are grizzled gold to greenish, limbs are grayish, and underside is white. Tip of the tail is creamy-white, and there is a small tuft of white fur at the base. Face is black. There is a pronounced but sinuous and tapered white band on the forehead, and long; stiff, yellowish (with black tips) cheek whiskers, the latter separated from the forehead band by a black stripe running from the corner of each eye back along temples. Scrotum is sky blue and surrounded by a zone of long orange hairs. Hairs on the Tantalus Monkey's hands and feet are not as dark as seen in the Vervet Monkey (C. pygerythrus); they tend to be light to medium gray. Some Tantalus Monkeys in southern Nigeria have such light-colored hands that they appear whitish. Tantalus Monkeysliving in higher elevations have thick coats. In C. t. tantalus, dorsal color tendsto be olive-green. The “ Jebbel Marra Tantalus ” (C. t. marrensis) is generally a lighter olive-brown above than C. i. tantalus. “ Budgett’s Tantalus ” (C. t. budgetti) is olive-brown dorsally with darker hands, feet, and tail tip. Cheek whiskers are markedly yellow with black speckling.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFECFFEAFAF46697FB04FBD3.taxon	biology_ecology	Habitat. Savanna, open woodland, montane forest, and forest-grassland mosaic, especially close to rivers. The Tantalus Monkey is extremely adaptable, residing in degraded habitats or forest edges (e. g. Togo and Benin), and it is found in rural and urban environments. They have been recorded up to ¢. 1900 m above sea level.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFECFFEAFAF46697FB04FBD3.taxon	food_feeding	Food and Feeding. The Tantalus Monkey has an eclectic diet, feeding on leaves, fruits, seeds, flowers, gums, eggs, invertebrates, and some small vertebrates. Their preferences for each category of food type vary greatly by season and study site. For example, in two studies in Cameroon, 61 % of the diet was fruit in one study, while fruits made up only 26 - 8 % of the diet in the other. Their terrestrial habits, combined with a wide selection offorest edge and grassland species, make Tantalus Monkeys key seeddispersers, an important consideration in forest regeneration of their habitats. They potentially disperse seeds as far as 1 - 5 km from the parent tree. Tantalus Monkeys readily raid crops and will often take handouts from tourists.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFECFFEAFAF46697FB04FBD3.taxon	breeding	Breeding. Female Tantalus Monkeys do not exhibit a sexual swelling, and menstrual flow is often difficult to detect. Mating takes place over 3 — 4 days, and the gestation period is ¢. 6 months. Hairs of neonates are uniformly dark gray, almost black, yet sparse. They acquire their adult coloration at c. 4 - 5 months of age. As with other species of Chlorocebus (and many other genera of primates, in fact), the sex of a newborn is often difficult to assess; the clitoris can be relatively large and the labia have a blue tnt, reminiscent of testes. Proper identification can be made by ignoring these factors and focusing only on location (i. e. male genitalia are ventrally located and best ascertained when an infant stands bipedally).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFECFFEAFAF46697FB04FBD3.taxon	activity	Activity patterns. Tantalus Monkeys are diurnal and largely terrestrial. They spend c. 20 - 25 % of their time feeding, but this varies by sex, season, and location of study. Much of their feeding time is spent on the ground or relatively low in the trees. They tend to use the ground to move long distances rather than moving through trees.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFECFFEAFAF46697FB04FBD3.taxon	biology_ecology	Movements, Home range and Social organization. Tantalus Monkeys live in large groups, often more than 70 individuals. A large group can occupy a home range ofas large as 90 ha, but there is seasonal variation and ranges depend on dispersion and abundance of key food sources. Individuals can move 2500 m / day. There are typically more adult females than adult males in a group. Tantalus Monkeys live in sympatry with Patas Monkeys (Erythrocebus patas) in much of their distribution.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFECFFEAFAF46697FB04FBD3.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Least Concern on The IUCN Red List. The subspecies have not been assessed. The Tantalus Monkey islisted as Class B in the African Convention on the Conservation of Nature and Natural Resources. It is common and widely distributed in appropriate habitats, and there are no major threats. The population is stable and possibly increasing due to forest clearing, which may provide it with more suitable areas in which to live. The subspecies tantalus is known to occur in 22 protected areas: Pendjari and West of the Niger national parks in Benin; Bénoué, Bouba-Ndjida, Kalamaloué, and Waza national parks, Faro Reserve, and Kimbe River and Mbi Crater game reserves in Cameroon; West of the Niger National Park in Niger; Cross River, Gashaka-Gumti, and Kainji Lake national parks, Oban Hills and Olokemeji forest reserves, and Borgu, Kwiambana, Pandam, Upper Ogun, and Yankari game reserves in Nigeria; and Keran and Malfacassa reserves in Togo. Budgett’s Tantalus occurs in Murchison Falls and Ruwenzori national parks and Budongo and Kasokwa forest reserves in Uganda. The Jebbel Marra Tantalus is not found in any protected areas.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEFFFEAFA2E6113F75EF532.taxon	materials_examined	Democratic Republic of the Congo, Lower Congo, Banana, fixed by E. Schwarz in 1928.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEFFFEAFA2E6113F75EF532.taxon	discussion	C. cynosuros is most often listed as a subspecies of C. aethiops, but is considered a subspecies of C. pygerythrus by J. Kingdon in 1997, and was separated as a full species by C. P. Groves in 2001. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEFFFEAFA2E6113F75EF532.taxon	distribution	Distribution. S Gabon, S Republic of the Congo, S DR Congo, Angola, Zambia (W of the Luangwa Valley), N Namibia, and N Botswana.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEFFFEAFA2E6113F75EF532.taxon	description	Descriptive notes. Head-body 34 - 65 cm (males) and 30 - 50 cm (females), tail 46 — 76 cm (males) and 35 - 66 cm (females); weight 3.1 - 6.4 kg (males) and 1.5 - 4.9 kg (females). The Malbrouck Monkey is similar to the Vervet Monkey (C. pygerythrus), but it is paler olive-gray. It is the only species of Chlorocebus with a light colored face (fleshypink with dark blotches). Bare skin of palms and soles is also pale, and callosities are pink. Cheek whiskers are long and directed upward and backward. Females resemble males but are smaller. Scrotum is a rich lapis blue.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEFFFEAFA2E6113F75EF532.taxon	biology_ecology	Habitat. Savanna, woodland, and forest edge. The Malbrouck Monkey is usually found in close proximity to rivers. It is a highly adaptable species and may be found in rural and urban environments, adjusting easily to human presence.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEFFFEAFA2E6113F75EF532.taxon	food_feeding	Food and Feeding. The Malbrouck Monkey feeds on fruit, seeds, flowers, gum, and bird eggs. Invertebrates and some small vertebrates (e. g. fledglings and lizards) are also included the diet, but detailed dietary data are not available.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEFFFEAFA2E6113F75EF532.taxon	breeding	Breeding. Breeding habits of the Malbrouck Monkey are similar to those of the Grivet Monkey (C. aethiops), with a single offspring born after a gestation of ¢. 6 months. Neonates are dark in color.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEFFFEAFA2E6113F75EF532.taxon	activity	Activity patterns. Malbrouck Monkeys are diurnal and mainly terrestrial.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEFFFEAFA2E6113F75EF532.taxon	biology_ecology	Movements, Home range and Social organization. The Malbrouck Monkey lives in large groups of up to 50 individuals. There is usually a balanced ratio of males to females. Long considered a subspecies of the Grivet Monkey, most observations suggest very similar social organization and ranging patterns.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEFFFEAFA2E6113F75EF532.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Least Concern on The [UCN Red List. The Malbrouck Monkey is listed as Class B in the African Convention on the Conservation of Nature and Natural Resources. It is wide-ranging and common, because much of the area where it occurs has a low human density. There are no major threats. It occurs in at least three protected areas: Luando Special Reserve in Angola, Chobe National Park in Botswana, and Etosha National Park in Namibia.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEFFFEAFF246167F782FC46.taxon	materials_examined	“ Cape Verde Islands, ” probably Senegal.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEFFFEAFF246167F782FC46.taxon	discussion	There may be some limited interbreeding of C. sabaeus with C. tantalus in Ghana. This species was long considered part of the genus Cercopithecus and often treated as a subspecies of C. aethiops. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEFFFEAFF246167F782FC46.taxon	distribution	Distribution. From Senegal and Guinea-Bissau (including the Bijagos Archipelago), E to just W of the Volta River system in Ghana; possibly also in S Mauritania. Introduced into the Cape Verde and Canary Is, as well as to Barbados, Nevis, and Saint Kitts Is in the West Indies.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEFFFEAFF246167F782FC46.taxon	description	Descriptive notes. Head — body 42 - 60 cm (males) and 30 - 50 cm (females), tail 46 - 76 cm (males) and 41 - 66 cm (females); weight 3.1 - 6.4 kg (males) and 1 - 5 — 4 - 9 kg (females). The Green Monkey is grizzled golden-green above, the color extending to limbs and base of tail. Undersurfaces are creamy white, backs of thighs are yellow, hands and feet are gray, and tail tip is golden-yellow. Face is black, with long yellow cheek whiskers, which are directed upward in front of ears and over temples from a whorl just in front of the relatively large (usually black) ears, and sharply distinct from the speckledgreenish crown. There is sometimes (but not always) a light brow band, much less noticeable than in other species of Chlorocebus. Scrotum is very pale blue. Legs are longer than arms. At birth, fur is dark, with face, ears, palms, and soles unpigmented (i. e. light colored). As a youngster ages, fur changes color well before that of the skin. Infants’ tails are relatively shorter than those of adults, and limbs are relatively longer.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEFFFEAFF246167F782FC46.taxon	biology_ecology	Habitat. Broad range of habitats, often preferring riparian gallery woodland, Acacia savanna, coastal scrub, or lowland tropical moist forest edges. Green Monkeys adapt well to living in mangrove swamps; they are often found in deltas of West African rivers. They are also common in riparian forest galleries and may frequent cultivated areas.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEFFFEAFF246167F782FC46.taxon	food_feeding	Food and Feeding. Green Monkeys eat a wide variety of leaves, fruits, and animal prey. They also eat gum from thorn trees (Acacia, Fabaceae), flowers, and seeds. Groups living near the sea often catch and eat crabs on mudflats at low tide; one population living in a mangrove swamp in Senegal fed largely upon fiddler crabs. As expected, their diet varies from month to month and from place to place, and there is great variation in feeding patterns among study sites. These differences are a direct result of food availability and dispersion of plants. Green Monkeys raid crops and will take handouts from tourists.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEFFFEAFF246167F782FC46.taxon	breeding	Breeding. Female Green Monkeys do not have sexual swellings. In Niokolo-Koba, the birth season is at the height of the dry season in January-March. In contrast, in dry habitat of northern Senegal, births are concentrated in the wet season (June — September). First parturition can occur as early as 2 - 5 years of age, although some first births have been reported at 4 - 5 years. The interbirth interval is c. 12 months. The gestation period is ¢. 6 months. Weaning can begin when an infant is as young as three months.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEFFFEAFF246167F782FC46.taxon	activity	Activity patterns. Green Monkeys are diurnal and mainly terrestrial. They may spend as much as one-third of their time on the ground. They often sleep in gallery forest in dry habitats but may range outside such areas by day. Studies of activity patterns for the Green Monkey provide conflicting data, likely because of differences in habitat type and data collection methods. Although feeding and moving activities vary greatly between study sites, one relatively stable figure is 7 - 9 % of the day spent socializing.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEFFFEAFF246167F782FC46.taxon	biology_ecology	Movements, Home range and Social organization. Groups of Green Monkeys often contain two or more adult males. Group size varies; most studies have found 18 - 33 individuals / group, but a study in Senegal estimated 174 individuals in one group. Daily movements are 665 - 2670 m, and home range can exceed 138 ha. Female Green Monkeys remain in their natal group, while males disperse. Aggressive behavior is relatively rare, but it may occur when one male is attacking another in the group or when two groups compete for access to a food resource. The latter event will often be accompanied by males leaping about in the trees while displaying their canines and erect penises. Females may participate in “ chuttering calls ” during this male display. Both sexes will lead departures from trees. Population densities are variously estimated at 1 - 2 — 4 - 4 ind / km?.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEFFFEAFF246167F782FC46.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Least Concern on The [UCN Red List. The Green Monkeyislisted as Class B in the African Convention on the Conservation of Nature and Natural Resources. It is widespread and reasonably abundant. There are no known major threats, and thus population numbers appear to be stable. Nevertheless, they are hunted for meat in Senegal. The Green Monkeys’ adaptability to a wide range of habitats often places them in contact with farmland. They are considered a pest in much of their distribution and can face harm or death from farmers protecting their crops. The Green Monkey occurs in three protected areas: Abuko Nature Reserve in Gambia and Basse Casamance and Niokolo-Koba national parks in Senegal.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEFFFEBFA2C6E07F71EFC6D.taxon	materials_examined	Africa (= South Africa).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEFFFEBFA2C6E07F71EFC6D.taxon	discussion	Further taxonomic work is required to assess the validity of the proposed subspecies. C. pygerythrus hybridizes with C. ae thiops in south-western Ethiopia and with C. tantalus in Uganda and in DR Congo. In northern Tanzania, some C. pygerythrus have very pale hands and feet, leading to questions about the descriptive features of this species. Dwarf forms (nesiotes and excubitor) are found on islands in the Indian Ocean and Lake Victoria. Five subspecies recognized.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEFFFEBFA2C6E07F71EFC6D.taxon	distribution	Subspecies and Distribution. C. p. pygerythrusF. Cuvier, 1821 — Botswana, Zimbabwe, SMozambique, Namibia, SouthAfrica (Kwazulu-NatalandTransvaalprovinces), Swaziland, andLesotho, SoftheZambeziRiver. C. p. excubitorSchwarz, 1926 — NEKenya (PateandMandaIsintheLamuArhipelago). C. p. hilgertiNeumann, 1902 — SEthiopia (EthiopianRiftValleyandintopartsofthehighlandsEoftheRift), SWSomalia, EUganda, Kenya, andNTanzania. C. p. nesiotesSchwarz, 1926 — ETanzania (PembaandMafiaIs). C. p. rufoviridis 1. Geoffroy Saint-Hilaire, 1843 — SW Uganda (S of Lake Victoria), Rwanda, Burundi, W Tanzania, Malawi, N Mozambique, and E Zambia (E of the Luangwa Valley) to the Zambezi River.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEFFFEBFA2C6E07F71EFC6D.taxon	description	Descriptive notes. Head-body 42 - 70 cm (males) and 30 - 62 cm (females), tail 45 - 76 cm (males) and 41 - 66 cm (females); weight 3.1 - 6.4 kg (males) and 1.5 - 4.9 kg (females). The Vervet Monkeyis grizzled-gray or olive above. This color extends to outer surfaces of limbs (which are not gray as they are in the Tantalus Monkey, C. tantalus) and is darker (often black) on hands, feet, and tail tip. Cheek whiskers are very short and combine with a white brow band to completely encircle the otherwise black face, with the former grading into the greenish speckled crown and neck. Fur at base oftail and in perineal region is bright red and does not form tufts, as it does in most Tantalus Monkeys. Scrotum is turquoise-blue; this color has been found to vary in some study subjects, perhaps related to stress, nutrition, or dominance status. The “ Black-chinned Vervet Monkey ” (C. p. pygerythrus) is ashy-gray to olive-green above, tending to be grayer in the west and more greenish in the east. “ Hilgert’s Vervet Monkey ” (C. p. hilgerti) is generally a paler brownish-yellow than other subspecies, often lacking the dark feet and hands of the other subspecies. The “ Wittu Islands Vervet Monkey ” (C. p. excubitor) resembles Hilgert’s Vervet Monkey, but is smaller. The “ Reddish-Green Vervet Monkey ” (C. p. rufoviridis) is a large subspecies, with orangey-yellow or fawn upperparts and (often) a reddish-infused underside. Cheek whiskers are long and speckled. A form from the Mahale Mountains region in western Tanzania is geographically close to the Reddish-Green Vervet Monkey, but lacks the reddish or orange tint that at least some Reddish-Green Vervet Monkeys have, and phenotypically is difficult to ascribe to a subspecies. The “ Pemba Vervet Monkey ” (C. p. nesiotes) resembles the Reddish-Green Vervet Monkey, but is smaller.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEFFFEBFA2C6E07F71EFC6D.taxon	biology_ecology	Habitat. Savanna, open woodland, forest-grassland mosaic, forest edge, cultivated areas, and highland regions to 1800 m above sea level. Vervet Monkeys are often associated with miombo woodland. They are generally absent from desert areas and deep forest. Vervet Monkeys in game parks have adapted well to tourist activities and are pests around some lodges.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEFFFEBFA2C6E07F71EFC6D.taxon	food_feeding	Food and Feeding. Vervet Monkeys have a diverse diet and feed on leaves, fruits, seeds, gums, flowers, and small animal prey. They raid crops and orchards and readily acclimate to humans.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEFFFEBFA2C6E07F71EFC6D.taxon	breeding	Breeding. Female Vervet Monkeys reproducefor the first time at 4 - 6 years of age. They may reproduce yearly, but in most habitats with reduced food availability, they reproduce at a slower rate. Phytochemical contents of some ingested flower species have been implicated in influencing the reproductive cycles of Vervet Monkeys. There is no sexual swelling in the female. Gestation lasts c. 6 months, followed by a single birth. The neonatal hair is dark, gradually turning to the adult coloration by six months of age. The face is pale at birth, turning darker gradually over the first few months.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEFFFEBFA2C6E07F71EFC6D.taxon	activity	Activity patterns. Vervet Monkeys are diurnal and mainly terrestrial. In general, they spend 30 - 40 % of their time feeding and a similar amount of time scanning for predators and conspecifics. These figures are subject to variation based on season, locale, and a variety of other factors. Most importantly, the effect of group size on activity patterns provides clear evidence that Vervet Monkeys derive benefits from living in larger groups.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEFFFEBFA2C6E07F71EFC6D.taxon	biology_ecology	Movements, Home range and Social organization. Vervet Monkeys live in multimale — multifemale groups of up to 75 individuals. Many unrelated males can be present in a group at any given time. Males transfer among groups, whereas females remain in their natal group (philopatric) and maintain fairly strict matrilineal relationships. This provides ample opportunity for allomaternal care and complex relationships among siblings — social behaviors that are also seen in other species of Chlorocebus. Often aggressively territorial, Vervet Monkeys have been observed to shift their home ranges away from other groups. On other occasions, they have moved home ranges in response to habitat deterioration. Such shifts have been observed to result in group fusion in some cases. The home rangesize is ¢. 23 ha, but varies by location and habitat type. Although considered a “ savanna species, ” their home ranges are typically restricted to areas with an ample number of trees. The vocal repertoire of Vervet Monkeys has been well studied. It includes a variety of predator-specific alarm calls that are used to communicate danger to conspecifics. Vervet Monkeys are favored prey of Leopards (Panthera pardus) in some regions. Pythons and martial eagles (Polemaetus bellicosus) also prey on them to a lesser extent.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEFFFEBFA2C6E07F71EFC6D.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Least Concern on The IUCN Red List. The subspecies have not been assessed. The Vervet Monkeyis listed as Class B in the African Convention on the Conservation of Nature and Natural Resources. It is widely distributed and often common; however, it is very patchily distributed, probably because of its need to drink water daily. There are no major threats. Vervet Monkeys are considered vermin in parts of their distribution, and they are actively persecuted (shot and hunted) by landowners in areas where they raid crops or interact with humans. They are also hunted for their meat in some areas (e. g. the Turkana district of Kenya). Vervet Monkeys occur in most protected areas in their range. The Black-chinned Vervet Monkey occurs in at least eleven protected areas: Maputo Game Reserve in Mozambique; Addo Elephant, Kruger, Mountain Zebra, Richtersveld, and Tsitsikamma national parks, Burman Bush and Oribi Gorge nature reserves, and Ndumu and Hluhluwe-Umfolozi game reserves in South Africa; and Zimbabwe National Park in Zimbabwe. Hilgert’s Vervet Monkey occurs in Samburu and Amboseli national parks in Kenya and Lake Manyara and Serengeti national parks in Tanzania. The Pemba Vervet Monkey occurs in Ngezi Forest Reserve in Tanzania. The Reddish-Green Vervet Monkey occurs in Gombe Stream, Mahale Mountains, Mikumi, Ruaha, and Udzungwa Mountains national parks in Tanzania and Lower Zambezi National Park in Zambia. The Wittu Islands Vervet Monkey is not found in any protected areas. A lack of updated records that take into account this most recent classification of subspecies of Vervet Monkeys suggests that this list may need careful reassessment.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEEFFF4FAFD66C9FBC2FDD6.taxon	materials_examined	Ethiopia, bamboo forest near Abera, east of Lake Abaya, 3300 m.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEEFFF4FAFD66C9FBC2FDD6.taxon	discussion	C. djamdjamensis was formerly listed as a subspecies of C. aethiops and has also been considered a species of the genus Cercopithecus. There is photographic evidence to suggest that hybridization occurs with C. aethiops on the margins of its distribution. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEEFFF4FAFD66C9FBC2FDD6.taxon	distribution	Distribution. W Ethiopia, in the Bale Mts, E of the Rift Valley lakes of Abiata, Shalla, and Zway.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEEFFF4FAFD66C9FBC2FDD6.taxon	description	Descriptive notes. Head-body 42 - 60 cm (males) and 30 - 50 cm (females), tail 46 76 cm (males) and 41 - 66 cm (females); weight 3.1 - 6.4 kg (males) and 1 - 5 — 4 - 9 kg (females). The Bale Mountains Monkey is a relatively short-tailed species, with long thick fur. Its upper body is deep grizzled-brown, with a creamy-white underside (mixed with gray on chest), grayish limbs, and dark gray hands and feet. Tail is grayish, with an inconspicuous red-brown tuft at its base. Face is black except for a short but very bushy white beard and cheek ruff, and a fine white moustache. A vague white brow band is separated from the cheek ruff by a broad black band running from eye to ear. Scrotum is blue.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEEFFF4FAFD66C9FBC2FDD6.taxon	biology_ecology	Habitat. High-elevation bamboo forest and forest edges at 2400 - 3000 m above sea level. Although previously described as living only in bamboo forests, new evidence suggests the Bale Mountains Monkey is more flexible in its habitat choice. Nevertheless, it does appear to have rather specialized and unusual habitat requirements, unlike other species of Chlorocebus.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEEFFF4FAFD66C9FBC2FDD6.taxon	food_feeding	Food and Feeding. The first behavioral and ecological study on the Bale Mountains Monkey was conducted as recently as 2007 - 2008. They eat primarily young leaves (80 - 2 % in one study). They also eat fruits, seeds, and animal prey (including invertebrates, reptiles, birds, and small mammals). Unlike other species of Chlorocebus, they appear to be specialized for eating bamboo leaves.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEEFFF4FAFD66C9FBC2FDD6.taxon	breeding	Breeding. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEEFFF4FAFD66C9FBC2FDD6.taxon	activity	Activity patterns. The Bale Mountains Monkey is diurnal and somewhat less terrestrial than other species in Chlorocebus. Feeding accounts for 65 - 7 % of their daily activity budget, with the remaining time spent moving (14 - 4 %), resting (10 - 7 %), and socializing (7 - 1 %).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEEFFF4FAFD66C9FBC2FDD6.taxon	biology_ecology	Movements, Home range and Social organization. The Bale Mountains Monkey lives in groups of up to 50 - 60 individuals. Their daily movement averages 928 m, and the entire home range is c. 15 ha. These figures come from two neighboring groups in one short study. Data indicated that the larger group traveled further each day and occupied a larger home range — a pattern suggesting that scramble competition may have occurred. Vocalizations of the Bale Mountains Monkey are similar to those of the Grivet Monkey (C. aethiops).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFEEFFF4FAFD66C9FBC2FDD6.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Vulnerable on The IUCN Red List. The Bale Mountains Monkey is listed as Class B in the African Convention on the Conservation of Nature and Natural Resources. It was presumably more widespread and abundant in historic times. It occurs at low densities and has recently been found in multiple forest fragments in southern Ethiopia, thus extending its distribution to the Guji and West Arsi zones of the Oromia Region and the Sidama Zone of the Southern Nations Nationalities and Peoples Region. Its main threats are ongoing habitat loss, fragmentation, and degradation. Harenna Forest in the Bale Mountains, where individuals are occasionally found, is under threat from encroaching human populations, fire, agriculture, and the removal of forest products. Of special concern is the removal of bamboo, a key resource for the Bale Mountains Monkey. Persecution for crop raiding may also be a problem. Bale Mountains Monkeys may be locally common in some areas (e. g. Odobullu Forest at 6 ° 87 ° N, 40 ° 17 ° E). Survey work in the Bale Mountains may eventually provide more information about its distribution and population status. It is known to occur in only one protected area, Bale Mountains National Park in Ethiopia, but it is also found in the proposed Harena-Kokosa National Forest Reserve. Although the Bale Mountains Monkey was first named in 1902, it has not been well studied. Until 2008, it waslisted as Data Deficient by ITUCN. This lack of information hampers our understanding of this species and prevents proper conservation action. More detailed studies are needed.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF1FFF4FF286627F714FB0E.taxon	materials_examined	Congo (= DR Congo). Restricted by R. Pocock in 1907 to Tschepo River, near Stanleyville (= Kisangani).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF1FFF4FF286627F714FB0E.taxon	discussion	A. lhoesti 1 s sympatric with Cercopithecus hamlyn: and C. mitis, in their respective distributions. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF1FFF4FF286627F714FB0E.taxon	distribution	Distribution. E DR Congo (from the Lualaba River E to the Ituri Forest, with an additional population S of the lower Lindi River), SW Uganda, Rwanda, and N Burundi.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF1FFF4FF286627F714FB0E.taxon	description	Descriptive notes. Head-body 54 - 70 cm (males) and 45 - 55 cm (females), tail 50 - 76 cm (males) and 46 - 67 cm (females); weight 6 - 10 kg (males) and 3.4 - 5 kg (females). Coat of L'Hoest’s Monkey is dense and black (there is some gray ticking on crown and nape), and limbs, flanks, and underside are also black. There is a short, oval-shaped, orangespeckled saddle on the lower back that stops short of the tail base. Except for a dark muzzle, face is gray; lighter coloring surrounds eyes and cheeks. These hairs are gray at the base, with four pairs of white and black bands. This differs from Preuss’s Monkey (A. preussi), which has only two pairs ofalternating black and white bands. L’ " Hoest’s Monkey has bushy white cheek whiskers and a white throat ruff. Ears are visible. Tail is gray above exceptfor the terminal one-fourth, which is black and tufted, and its ventral surface tends to be light in color. Tail of L’Hoest’s Monkey is sometimes said to be “ partially prehensile, ” but this belief is based on the fact that it is normally carried with a pronounced hook nearthe tip. Feet are narrow and well adapted for running on the ground. Several features of the anatomy of L’Hoest’s Monkey are associated with terrestrial adaptations. Scrotum is light blue, and females lack a sexual swelling.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF1FFF4FF286627F714FB0E.taxon	biology_ecology	Habitat. A variety of forest types, including lowland, riparian, submontane, montane, and dry woodland forests. ' Hoest’s Monkeys are occasionally found in cultivated land. They normally live at elevations of ¢. 1000 - 2500 m, although they have been recorded up to 2900 m. Forthis reason, L'Hoest’s Monkeys, and closely related Preuss’s Monkeys, are often called “ Mountain Monkeys. ” Because they are largely terrestrial, they eat many fruits and seeds from shrubs and secondary tree species and thus tend to be generalists in their use of habitat types.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF1FFF4FF286627F714FB0E.taxon	food_feeding	Food and Feeding. I. ’ Hoest’s Monkeys are omnivorous, eating terrestrial herbs, fruits, seeds, young leaves, flowers, berries, insects, bird eggs, wood, lichens, sap, and arthropods. About 25 - 45 % of their waking hours are spent feeding and foraging (this varies by location). In a study in Nyungwe Forest National Park, Rwanda, terrestrial herbaceous vegetation made up more than 35 % ofthe diet, with fruits comprising only 24 %. The top three food items (50 % of the diet) were terrestrial herbs (several species) and seeds of Macaranga neomildbraediana and Alchornea hirtella (both Euphorbiaceae). In contrast, a population studied in Kalinzu National Park, Uganda, spent 66 % ofits feeding time on invertebrates. At both sites, L'Hoest’s Monkeys ate more invertebrate species in the dry season than at other times of the year. These findings underscore the broad environmental adaptability of L’Hoest’s Monkey. Unfortunately, they occasionally raid crops from nearby villages, thus placing them at risk from retaliation from humans.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF1FFF4FF286627F714FB0E.taxon	breeding	Breeding. Reproductive activities of L’Hoest’s Monkeys are seasonal. Births tend to occur at the end of dry season (November — April). After a gestation of ¢. 5 months, a single infant is usually born at night. Newborns are born uniformly brown, without the ticking pattern. They acquire adult coloration over the first 2 - 3 months oflife. The interbirth interval is c. 2 years.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF1FFF4FF286627F714FB0E.taxon	activity	Activity patterns. [. ' Hoest’s Monkeys are diurnal and mostly terrestrial. They make limited use of arboreal space for foraging, avoiding predators, and sleeping. They choose a wide variety of sleeping sites compared with sympatric Blue Monkeys (Cercopithecus mitis). A study group in Rwanda spent 38 % ofits time on the ground and only 27 % in upper tree canopies. They spend an average of 45 % of their time during the day foraging and feeding.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF1FFF4FF286627F714FB0E.taxon	biology_ecology	Movements, Home range and Social organization. Group sizes of I. Hoest’s Monkeys are usually 10 - 17 individuals, although a study group in Nyungwe Forest National Park, Rwanda, numbered 29. Groups typically include only one mature male, several adult females, and their offspring. In the Rwandan study, the home range was 116 - 6 ha. The monthly travel distance covered 24 - 70 % of the home range. As expected, ranging patterns were altered by changes in availability of fruit sources, but there was not a significant seasonal influence, and rainfall was not related to travel distance in the Rwandan study. Groups are territorial; intergroup encounters usually involve chasing and vocalizing, often lasting for more than one hour.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF1FFF4FF286627F714FB0E.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Vulnerable on The [UCN Red Last (as Cercopithecus lhoesti). I’ Hoest’s Monkey is listed as a Class B species of the African Convention on the Conservation of Nature and Natural Resources and protected by national legislation in Burundi, Rwanda, and Uganda. It is threatened by deforestation on the eastern edge ofits distribution. This habitat destruction is mainly a consequence of agricultural expansion, resulting in frequent opportunities for crop raiding and other conflict with people. L'Hoest’s Monkey is also hunted for its meat and fur in some areas. Its relatively large size and terrestrial habits make it vulnerable to snaring and shotgun hunting. It is also an important prey for Leopards (Panthera pardus) in the Ituri Forest. It is known to occurin at least eleven protected areas: Kibira National Park in Burundi; Kahuzi-Biéga and Virunga national parks, Okapi Wildlife Reserve, and Tayna Gorilla Reserve in DR Congo; Nyungwe Forest National Park in Rwanda; and Bwindi Impenetrable, Kibale Forest, and Ruwenzori national parks and Kalinzu and Maramagambo forest reserves in Uganda. Because the entire distribution of L'Hoest’s Monkey is currently in a region of political and military conflict, declining populations are expected to continue. Given a 10 year generation time, the overall population reduction may exceed 40 % over a 30 year timeframe. Controlling bushmeat hunting and improved land and water management are among the recommended conservation actions.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF1FFF5FA32602BFCE6F90F.taxon	vernacular_names	Preuss’s 9 Monkey	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF1FFF5FA32602BFCE6F90F.taxon	materials_examined	Cameroon, Victoria.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF1FFF5FA32602BFCE6F90F.taxon	discussion	Two subspecies are recognized.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF1FFF5FA32602BFCE6F90F.taxon	distribution	Subspecies and Distribution. A. p. preussi Matschie, 1898 — SE Nigeria and SW Cameroon highlands (Cross River Highlands, Korup National Park, Bamenda Highlands, and Mount Cameroon). A. p. insularis Thomas, 1910 — Bioko I (Equatorial Guinea).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF1FFF5FA32602BFCE6F90F.taxon	description	Descriptive notes. Head — body 42 - 70 cm (males) and 37 - 55 cm (females), tail 50 - 76 cm (males) and 41 - 67 cm (females); weight 4.7 - 10 kg (males) and 2.9 - 4.3 kg (females). Preuss’s Monkey is similar to L'Hoest’s Monkey (A. lhoesti), but it has only a small amount of white fur on its upper chest and throat, merging upward with gray cheek whiskers. Hairs on crown and nape are black, ticked with gray. It has a saddle ofsolid chestnutred. Flanks are gray with buff banding, underside is dark gray, and limbs are entirely black. Although tail is like the back proximally, it grades into a dark gray, with its terminal one-fourth being black. Preuss’s Monkey usually holds its tail with a hook or loop at the end, often described as a question mark. Face is uniformly gray, and ears are hidden by crown hairs. Eyes appear very dark reddish-brown. Gray hairs have only two pairs of white and black bands (unlike I'Hoest’s Monkey, which has four pairs). Scrotum is light blue. Female Preuss’s Monkeysare similar to males, but slightly smaller. The “ Cameroon Preuss’s Monkey ” (A. p. preussi) is the larger of the two subspecies, with a tail that is longer than head-body length. Dorsal saddle is a brighter chestnut than in the “ Bioko Preuss’s Monkey ” (A. p. insularis), and it extends more posteriorly. Tail is a lighter gray. The Bioko Preuss’s Monkey is smaller than the Cameroon Preuss’s Monkey, with a darker gray tail that is relatively shorter (i. e. about equal to head-body length). Dorsal saddle is less bright and does not extend as far on to the tail.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF1FFF5FA32602BFCE6F90F.taxon	biology_ecology	Habitat. Primary and secondary submontane and montane forest, preferring elevations of 800 - 2500 m. The Cameroon Preuss’s Monkey is found in highland forests of the Cameroon-Nigeria border regions. It may also occur in isolated forest patches in mountainous grasslands. On Bioko Island, the Bioko Preuss’s Monkey shows a preference for Schefflera (Araliaceae) forest above 1500 m, but it is also found in lowland forest (e. g. at 450 m in the Caldera on Bioko Island).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF1FFF5FA32602BFCE6F90F.taxon	food_feeding	Food and Feeding. Although there has been little systematic study of their feeding behavior, Preuss’s Monkeys are known to eat fruits, seeds, leaves, flowers, small aquatic animals, and termites. In the study at Kilum, 52 % of the diet of the Cameroon Preuss’s Monkey was fruit and seeds. As with other primates with diets of largely seasonally available foods, varied diets likely have important influences on movements and behavioral habits. The terrestrial nature of Preuss’s Monkeys probably serves them well in their high-elevation habitats, where the terrain is often broken by rocky outcrops.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF1FFF5FA32602BFCE6F90F.taxon	breeding	Breeding. There is no specific information for this species, but Preuss’s Monkey is assumed to have similar patterns to those of the Sun-tailed Monkey (A. solatus).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF1FFF5FA32602BFCE6F90F.taxon	activity	Activity patterns. Preuss’s Monkey is diurnal and largely terrestrial, spending most of its ime on the ground or in lowerlevels of the forest. The degree of terrestriality varies by study site, which may indicate a difference between the subspecies. On Bioko Island, more than 60 % of observations of the Bioko Preuss’s Monkey were on the ground or in the shrub layer. In contrast, a study in Cameroon recorded the Cameroon Preuss’s Monkey spending most of its time in trees but traveling on the ground to cross open spaces.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF1FFF5FA32602BFCE6F90F.taxon	biology_ecology	Movements, Home range and Social organization. A number of factors contribute to the lack of information for Preuss’s Monkeys. They are rare and of a very shy and secretive nature. Moreover, they live in areas that are often difficult to reach because of physical constraints or political restrictions. Consequently, home range size is not yet known. A study in Cameroon found variation in the composition of groups on different days in the same location (at similar times of day). This suggested either a flexible grouping pattern or overlap of home ranges. Nevertheless, Preuss’s Monkeys are normally found in unimale-multifemale groups of 2 - 19 individuals. On occasion, more than one adult male may be seen in the group. Single males are often seen, but never single females. Unlike many other guenons, Preuss’s Monkey rarely interacts with other primates. At dusk, adult males give one loud call, which is answered by neighboring males as the groups settle down for the night. This call is apparently absent in the other species of Allochrocebus. Alarm calls take the form of chirps in females and “ hacks ” in males.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF1FFF5FA32602BFCE6F90F.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Endangered on The IUCN Red Lust (as Cercopithecus preussi), including both subspecies. Preuss’s Monkey is listed as a Class B species of the African Convention on the Conservation of Nature and Natural Resources. Populations have been severely affected by habitat destruction and bushmeat hunting. Cameroon’s forests, for example, have been severely degraded by cultivation, fire, and the collection of wood for fuel. As a result, little montane forest remains in the mainland part of its range, and this forest area is not protected. Preuss’s Monkey is highly susceptible to hunting because it is semi-terrestrial and relatively large. Hunting has led to a decline in its population across its distribution, but particularly on Bioko Island where the estimated decline exceeds 50 % over a 20 year period. Recent reports indicate a steady increase in the number of carcasses of Bioko Preuss’s Monkeys found in the bushmeat market of Malabo on Bioko Island. In mainland Cameroon, Preuss’s Monkey is uncommon, and its populations are highly fragmented. Surveys of its status and distribution across its entire distribution are needed. Improved protection of montane forest, increased law enforcement, and banning of bushmeat hunting are urgently required. The Cameroon Preuss’s Monkey occurs in Takamanda, Mount Cameroon, and Ebo national parks in Cameroon and the Okwangwo Division of Cross River National Park and on the still unprotected Obudu Plateau in Nigeria. The Bioko Preuss’s Monkey occurs in Basilé National Park and Southern Highlands Scientific Reserve in Equatorial Guinea.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF0FFF5FFEA622CF90AF5D1.taxon	materials_examined	Gabon, River Bali, Foret des Abeilles, south-east of Booué (0 ° 14 ° S, 12 ° 15 ’ E).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF0FFF5FFEA622CF90AF5D1.taxon	discussion	This species is monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF0FFF5FFEA622CF90AF5D1.taxon	distribution	Distribution. C Gabon, S of the Ogooué River and W of the Offoué River, the W limit remains unclear, but it has been recorded E of the Waka National Park, the exact S limit is also unknown, butit extends to the foothills of the Massif du Chaillu. S limits may in fact be related to the presence of mountainous terrain typically avoided by this lowland forest species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF0FFF5FFEA622CF90AF5D1.taxon	description	Descriptive notes. Head-body ¢. 60 - 70 cm (males) and ¢. 50 - 55 cm (females), tail 65 - 76 cm (males) and 60 - 67 cm (females); weight 6 - 9 kg (males) and 3 - 5 — 4 - 5 kg (females). Crown and nape of the Sun-tailed Monkey are dark gray to black, with chestnut-orange speckling, and saddle is chestnut-orange, extending to flanks, shoulders, and upper thighs. Remainder of limbs is black, while upper chest and distinctive throat ruff are notably white. Tail is bright orange-yellow toward tip, and the remainderis whitish ventrally and speckled-gray dorsally. Similar to L'Hoest’s Monkey (A. lhoesti), the Sun-tailed Monkey carries its tail held horizontally and curving downward, and then bent upward with the tip falling back down. Face is dark, with a black muzzle and agouti-gray cheek whiskers. Ears are not hidden in the crown. Scrotum is powder blue.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF0FFF5FFEA622CF90AF5D1.taxon	biology_ecology	Habitat. Hilly moist evergreen forests, often bisected by rivers and streams. The Suntailed Monkey prefers densely shaded, tangled areas. It is a lowland forest species, with its natural habitat dominated by Gabon mahogany (Aucoumea, Burseraceae), the leguminous Awoura (Julbernardia, Fabaceae), false nutmeg (Pycnanthus, Myristicaceae), and Mubala-oil tree (Pentaclethra, Fabaceae).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF0FFF5FFEA622CF90AF5D1.taxon	food_feeding	Food and Feeding. The diet of the Sun-tailed Monkey is currently being studied, but it is known to contain fruits that are abundant throughout the year. It also eats seeds, grasses, small vertebrates, and insects. It occasionally raids crops from localvillages.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF0FFF5FFEA622CF90AF5D1.taxon	breeding	Breeding. Thefirst recorded birth of a Sun-tailed Monkey in captivity occurred in the early morning hours. At three days old, the infant weighed 382 g. The fur of infant Sun-tailed Monkeys is orange-streaked black, and their skin has little pigmentation. By eleven weeks of age, adult coloration begins to appear for both face and body. The interbirth intervalis just under two years.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF0FFF5FFEA622CF90AF5D1.taxon	activity	Activity patterns. The Sun-tailed Monkey is diurnal and semi-terrestrial. As a relatively newly discovered species, its daily activity patterns are still unknown.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF0FFF5FFEA622CF90AF5D1.taxon	biology_ecology	Movements, Home range and Social organization. Sun-tailed Monkeyslive in unimale — multifemale groups. On average, a group has c. 16 adult females. They appear to have few vocalizations and lack a loud call. It is thought that their tail coloring (white underside) and white throat ruff play a role in communication in their relatively dark forest habitat. They appearto thrive well in degraded secondary forests; i. e. population numbers remain stable after light logging because of the resulting increase in dense undergrowth. Population density varies with habitat suitability and seemingly elevation. At the Makandé Field Research Station, the density of Sun-tailed Monkeys was 25 ind / km?, but it was only c ¢. 4 ind / km? in southern Lopé National Park.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF0FFF5FFEA622CF90AF5D1.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Vulnerable on The IUCN Red List (as Cercopithecus solatus). The Sun-tailed Monkey is listed as Class B under the African Convention on the Conservation of Nature and Natural Resources. It was originally believed to be confined to its type locality in the Foret des Abeilles (south and west of the great bend of the Ogooué River and east of the Offoué River), but additional populations were discovered in 1994 and again in 2010, thereby extending the known distribution to ¢. 18,000 km? ®. The Sun-tailed Monkey was declared a “ totally protected species ” by the Gabonese government in 1994. Discovered in 1984, the Suntailed Monkey has become a flagship species for conservation in Gabon. Nevertheless, it continues to be hunted for its meat, despite being difficult to hunt because ofits secretive behavior. Its semi-terrestrial nature makes it vulnerable to the use of snares and dogs for hunting, butit is also killed by shotgun in growing numbers. The biggest threat to the Sun-tailed Monkey, however, is habitat loss, mainly due to logging; its type locality, the Forét des Abeilles, is largely allocated to logging concessions and is, therefore, unprotected. The Sun-tailed Monkey occurs in Lopé National Park in Gabon, and it has been seen very near the boundaries of Waka National Park and Birougou National Park. Only a small portion of the distribution of the Sun-tailed Monkey is in Lopé National Park, while the rest occurs in unprotected areas. Thus, better protection of its habitat is recommended. Monitoring of planned logging activity and better enforcement of hunting in the area are also urgently needed. Several Sun-tailed Monkeys are held in captivity in the Centre International de Recherches Médicales de Franceville in Gabon.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF5FFF0FFDA64A0F743FCBD.taxon	materials_examined	Congo-Zaire, Sankuru District, Lomela.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF5FFF0FFDA64A0F743FCBD.taxon	discussion	The original description of C. dryas was based on a photograph of a pet, 2 - year old juvenile male and its incomplete skin and skull in Lomela (2 ° 20 ° S, 23 ° 12 ’ E), a town on the Lomela River that marks the north-eastern boundary of Salonga National Park. The origin of this pet is unknown, but H. Schouteden argued, in 1946, that it may have been taken from Watsi, a village on the right bank of the Tshuape River, but the locality was never confirmed. In 1977, D. F. E. T. Van den Audenaerde described an adult guenon (believed to be male), from Wamba in the Zone de Djolu (¢. 250 - 260 km north of Lomela), as C. salongo. In 1991, M. Colyn, J. - P. Gautier, and Van den Audenaerde concluded that C. salongo is in fact an adult C. dryas, but E. Sarmiento, in 2002, argued that C. dryas and C. salongo may well, in fact, be distinct species and that further research was needed. In his note, Sarmiento indicated that, following the suggestion of Schouteden, there is a town called Watsi Kengo in the northern sector of Salonga National Park (c. 1 ° 00 ’ S, 20 ° 38 ’ E) where it might have come from. Watsi Kengo is a 1 - 2 day walk south from the fork of the Lomela and Tshuapa rivers, and there is a logic that the pet in the town of Lomela (upriver on the Lomela River) could have come from there. Djolu, near Wamba (0 ° 01 " N- 0 ° 01 " S, 22 ° 31 ’ - 33 " E), is the type locality of C. salongo. It is to the north of Watsi Kengo, south and west of the Lopori River. In his 1966 monograph, W. C. O. Hill concluded it was a subspecies of C. diana, and Thys van den Audenaerde in his description of C. salongo also suggested a close relationship with C. diana. The likeness is superficial, however, and aspects of their ecology and cranial morphology argue against it. The speciesis quite distinct, possibly a sister to all other Cercopithecus species, and with C. salongo as a junior synonym, C. P. Groves assigned C. dryas to its own species group, the dryas species group; this arrangement is followed here. Sarmiento did not discount the possibility that salongo may be a subspecies of C. dryas. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF5FFF0FFDA64A0F743FCBD.taxon	distribution	Distribution. Congo Basin in DR Congo (Equateur Province) in the Kokolopori Bonobo Reserve; it is unknown to people N of the Lopori. Its distribution is otherwise a mystery, but it is supposed it would extend between the Lopori River and the Luo River to the W; its possible occurrence in the Lolema River Basin to the S has not been confirmed.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF5FFF0FFDA64A0F743FCBD.taxon	description	Descriptive notes. Head-body 35 - 38 cm, tail 50 cm (females; no information for males); weight 3 kg (males) and 2 - 3 kg (females). The Dryas Monkey is one of the smallest guenons. Back and upper limbs are grayish chestnut. It has a creamy-white or white chin, and cheek-whiskers and a white brow, framing a triangular black face. White side whiskers are separated from the white brow band on either side by a thin posterio-superiorly directed bare, line of black facial skin. There are elongated tufts between crown and temples. Chest, inside upper arms and legs, and abdomen, extending to buttocks, are white. Adults have jet black lower limbs, both inside and out. A white patch on buttocks and posterior thighs extends along tail. Dark color on tail is restricted to a thin line along its upper surface and a black tip. The male hasslightly longer white fur on the chin than the female. Scrotum is light blue. In juveniles, black on limbs and tail is absent, and there is very little white on buttocks.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF5FFF0FFDA64A0F743FCBD.taxon	biology_ecology	Habitat. Mixed mature and secondary forest, but also tropical, riparian, and swamp forest to ¢. 380 m above sea level. Four small populations of Dryas Monkeys have been identified in mixed mature and young second growth forest in the Kokolopori Bonobo Reserve.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF5FFF0FFDA64A0F743FCBD.taxon	food_feeding	Food and Feeding. Dryas Monkeys feed mainly on fruits, supplemented with young leaves and shoots from plants such as Aframomum (Zingiberaceae) and Megaphrynium (Marantaceae).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF5FFF0FFDA64A0F743FCBD.taxon	breeding	Breeding. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF5FFF0FFDA64A0F743FCBD.taxon	activity	Activity patterns. Dryas Monkeys are diurnal and semi-terrestrial, generally using lower levels of the forest canopy. At Wamba, they are almost always reported on or close to the ground. They often forage on the ground. Locomotion is quadrupedal.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF5FFF0FFDA64A0F743FCBD.taxon	biology_ecology	Movements, Home range and Social organization. Dryas Monkeys are reported to live in multimale-multifemale groups of up to 30 individuals, but usually 2 - 15 individuals. They form mixed groups with other guenons such as Wolf’s Monkey (C. wolft) and the Red-tailed Monkey (C. ascanius) that occur in the same region. Little is known of Dryas Monkeys in their natural habitat.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF5FFF0FFDA64A0F743FCBD.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Critically Endangered on The IUCN Red List. The Dryas Monkey is listed as Class B in the African Convention on the Conservation of Nature and Natural Resources. Its known distribution is tiny, and itis clear from survey work that its numbers are low, probably not exceeding a few hundred individuals. All currently known individuals are present in the region along the left bank of the lower Lopori River, possibly restricted in the west by the Luo River. They are known to occur in just one protected area, Kokolopori Bonobo Reserve of 50,000 ha. There has been no recent information concerning their existence to the south in the Lomela River Basin. Reports to date have indicated that Dryas Monkeys occur near but not in Salonga National Park, but no surveys have been carried out specifically to address the possibility ofits occurrence in, for example, the northern sector. They are hunted for food. Conservation measures centered on the Dryas Monkey in the Kokolopori Bonobo Reserve are being carried out by the non-governmental organizations, Vie Sauvage and the Bonobo Conservation Initiative.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF5FFF1FA27669AFE2DF7AA.taxon	materials_examined	Guinea (probably Liberia).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF5FFF1FA27669AFE2DF7AA.taxon	discussion	C. diana and C. roloway comprise the diana species group. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF5FFF1FA27669AFE2DF7AA.taxon	distribution	Distribution. S Guinea (Forécariah Priovince), Sierra Leone, Liberia, and SW Ivory Coast (W of the Sassandra River).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF5FFF1FA27669AFE2DF7AA.taxon	description	Descriptive notes. Head-body 50 - 60 cm (males) and 42 - 45 cm (females), tail 85 cm (males) and 70 cm (females); weight 5 - 2 kg (males) and 3 - 9 kg (females). Together with the Roloway Monkey (C. roloway), the Diana Monkey is the largest of the forest guenons. Individuals are brightly colored, with strikingly patterned coats. Fur is speckled purple and gray, with a broad chestnut-brown saddle on lower back. Throat, chest, and anterior surfaces of upper arms are white, sharply demarcated from the dark color on other surfaces. Lower legs, belly, and tail are black, and buttocks and inner surfaces of thighs are a deep orange-red. An oblique white stripe runs along each outer thigh. Faceis black, and forehead is framed by a thin whitish crescent (with some reddish hairs intermixed). A short, square “ goatee ” hangs from the chin; it is white, fringed with black. Individuals raise their long tails in a high arching curve when moving.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF5FFF1FA27669AFE2DF7AA.taxon	biology_ecology	Habitat. Primary and old secondary lowland moist forest, gallery forest, and semideciduous forest near rivers. Diana Monkeys travel and forage mainly in the forest canopy. They are rarely found in degraded forest, but they are able to survive in lightly logged forest if the canopy remains.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF5FFF1FA27669AFE2DF7AA.taxon	food_feeding	Food and Feeding. The Diana Monkey is highly frugivorous; individuals studied in Tai Forest National Park spent 59 % oftheir time feeding on fruits. Commonly eaten fruits are those of the canopy trees Sacoglottis gabonensis (Humiriaceae) and Dialium aubrevillev (Fabaceae), besides those of the liana Landolphia hirsuta (Apocynaceae) and the stilt-rooted tree Uapaca guineensis (Euphorbiaceae). Flowers and leaves are seasonally important as a standby when fruits are scarce. Insects are also an important component of the diet, more so than in many other guenons, and they spend long periods carefully searching the foliage of canopy trees for insect prey.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF5FFF1FA27669AFE2DF7AA.taxon	breeding	Breeding. Although reproductive patterns of Diana Monkeys have not been comprehensively studied, young infants (light brown to gray in color) have been observed most frequently in the dry months ofJanuary-February on Tiwai Island in Sierra Leone, suggesting that they have a December — February birth peak. The gestation period is 5 — 6 months. Mothers usually give birth to a single infant and carry it ventrally, at first supported by hand. Females become sexually receptive about once a month and do not exhibit a genital swelling. Sexual maturity is most likely reached at 3 - 5 years for females and 4 - 5 - 6 years for males. Individuals may live more than 30 years; one captive individual lived forjust over 35 years.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF5FFF1FA27669AFE2DF7AA.taxon	activity	Activity patterns. Diana Monkeys are active, arboreal primates that spend most of their time in the middle and upper forest canopy. Forms of locomotion include walking and running (¢. 70 %), climbing quadrupedally (c. 20 %) through the forest canopy and leaping between canopy gaps (c. 10 %). Although the Diana Monkey usesall forest strata and occasionally moves on the ground, individuals spend most of their time above ground and use large boughs, especially for traveling, more than do the smaller guenons. As diurnal animals, groups spend the night in the topmost layer of the canopy, descending to the middle canopy by day to feed. Forty to 45 % of their active time is spent feeding and foraging and 25 % moving, although there is considerable variation in time allocation across groups, seasons, and years.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF5FFF1FA27669AFE2DF7AA.taxon	biology_ecology	Movements, Home range and Social organization. Movements, traveling distances, and home ranges of the Diana Monkey are reported to be highly variable and depend on factors such as group size, habitat features, and resource availability. The average distance traveled in a day is reported to be 1 - 1 - 5 km, and home ranges are 29 - 93 ha. Females and young chase their opponents in neighboring groups at territorial boundaries. Females also emit a “ chatter-scream ” that incites louder calls and displays by males in the group. Social groups tend to be relatively large, with 15 - 30 individuals. Typically, groups contain just one adult male, 5 - 13 adult females, and young. Solitary males that have not succeeded in joining a group move around on their own, occasionally joining groups of other species. Diana Monkeys are notably less affiliative with members of their group compared with other guenons. Males, in particular, exhibit low levels of integration into their group; females groom other females and young, but males are reported to never receive any grooming. Higher levels of agonistic interactions are also ascribed to the level of competition for food within groups. The main predators of Diana Monkeys are Leopards (Panthera pardus), Chimpanzees (Pan troglodytes), eagles, and humans. Research by K. Zuberbtuihler and colleagues, using playbacks of vocalizations of Leopards and crowned hawk-eagles (Stephanoaetus coronatus), showed that Diana Monkeys gave alarm calls that were specific for Leopards and different from those for eagles. They would often approach the source of the calls, letting other group members know of the danger and which predator had been seen. Leopards and eagles hunt by surprise, and alarm calls result in them calling off a hunt. The approach of humans and a playback of social screams of Chimpanzees, however, generally did not result in alarm calling. Both are pursuit predators, and the monkeys’ alarm calls would evidently draw their attention. Diana Monkeys instead keep quiet and try to hide and slip away unnoticed. Leopards are also predators of Chimpanzees, and Chimpanzee leopard-alarm screams (as opposed to social screams) stimulate bouts of Leopard alarm-calling by Diana Monkeys. Diana Monkeys travel with groups of Campbell's Monkeys (C. campbelli) and respond to their alarm calls, which are also predator specific. Although evidently serving as alarm calls for predators, the loud calls of Diana Monkeys differ between sexes, and males show a peak in loud calling in the early evening that is not related to predators. They have a slightly different structure (a different number ofsyllables), group members do not respond with their own calls, and loud calling in this situation is believed to play a role in the male’s defense of his females and territory.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF5FFF1FA27669AFE2DF7AA.taxon	conservation	Status and Conservation. CITES Appendix I. Classified as Vulnerable on The IUCN Red List. The Diana Monkey is listed as Class A in the African Convention on the Conservation of Nature and Natural Resources. Habitat loss and hunting have led to a 30 % decline in Diana Monkeys over the past 30 years (three generations). Habitat loss 1 s driven mainly by logging and clearing land for agriculture and charcoal production. Such deforestation continues to reduce habitat availability for Diana Monkeys. They are dependent on tall mature forest that is becoming increasingly degraded and fragmented. Diana Monkeys are also more vulnerable to hunting than other guenons because of their larger size, noisy vocalizations, and bright colors. They occur in a number of protected areas: Ta Forest National Park in Ivory Coast; Tiwai Island Wildlife Sanctuary and the proposed Gola and Loma national parks in Sierra Leone; and Sapo National Park in Liberia. Despite the protected status of these areas and official prohibition of hunting, Diana Monkeys are continuing to lose their habitat and are being harvested at unsustainable levels.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF4FFF1FFFF6D8FF809F5A4.taxon	materials_examined	Guinea (West Africa).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF4FFF1FFFF6D8FF809F5A4.taxon	discussion	Intermediate features between C. roloway and C. diana have been found near the Sassandra River, there the orange tones of the buttocks and inner thighs become deeper, but there is no intermediacy in the beard or brow band. C. rloway and C. diana comprise the diana species group. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF4FFF1FFFF6D8FF809F5A4.taxon	distribution	Distribution. SE Ivory Coast and SW Ghana, possibly Togo.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF4FFF1FFFF6D8FF809F5A4.taxon	description	Descriptive notes. Head-body 50 - 60 cm (males) and 42 - 45 cm (females), tail 85 cm (males) and 70 cm (females); weight 5 - 2 kg (males) and 3 - 9 kg (females). The Roloway Monkey closely resembles the Diana Monkey (C. diana) in appearance. Four main differences distinguish them: length and shape of beard (a much longer, pointed, and entirely white beard c. 75 mm in length on the Roloway Monkey); color of inner thighs and pubic region (creamy-white to orangey-yellow on the Roloway Monkey vs. orangered on the Diana Monkey); prominence of a band of white hair across brow (more prominent and extending further on sides on the Roloway Monkey); and thickness of the white thigh stripe (the Roloway Monkey's is narrower than Diana Monkey's).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF4FFF1FFFF6D8FF809F5A4.taxon	biology_ecology	Habitat. Primary and some secondary rainforest, gallery forest, and semi-deciduous forest near rivers. The Roloway Monkey has been much less intensively studied than the Diana Monkey, so fewer details of its habitat preferences (as well as ecology and behavior) are known, but they are presumed to have similar adaptations. The reported distribution of the Roloway Monkey also suggests that it may be able to occupy slightly drier and more seasonal forests than the Diana Monkey. The Roloway Monkey is a high-canopy specialist like the Diana Monkey.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF4FFF1FFFF6D8FF809F5A4.taxon	food_feeding	Food and Feeding. The only field study of the diet of the Roloway Monkey was done by S. Curtin in Bia National Park, western Ghana, in 1976 - 1977. The diet comprised pulp ofripe fruits (31 %), insects (25 %), seeds of ripe fruits (22 %), leaf buds (7 %), flower buds and flowers (6 %), and unripe fruit (5 %); seeds of unripe fruits, mature leaves, root nodes, and bulbs were eaten occasionally (less than 1 %). In the dry season, the pulp of mature fruit (especially of lianas and vines) was the most important food, followed by insects and the seeds of ripe fruits. In the wet season, ripe fruits and insects were equally important. During the study, seeds (large and black with a red aril and especially rich in oils) of Pycnanthus angolensis (Myristicaceae) and small immobile insects on branches and foliage, most particularly in the large legume Piptadeniastrum africanum (Fabaceae), were the most important food items in terms of the time dedicated to eating them. Both trees were common at Bia. Diets of Roloway and Diana monkeys can vary greatly among populations, years, and even neighboring groups. Diets of Roloway Monkeys at Bia contained 24 - 3 % ripe and unripe seeds (77 % of them from Pycnanthus and Funtumia elastica, Apocynaceae) in contrast to the diets of Diana Monkeys from three field studies that contained only 0 - 11 - 6 % seeds. Pycnanthus, like figs, produces fruits asynchronously, but it provides small quantities of fruits and seeds piecemeal over many months, unlike the mast fruiting of figs. The insect portion of the diet of Roloway Monkeys was similar to that of Diana Monkeys.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF4FFF1FFFF6D8FF809F5A4.taxon	breeding	Breeding. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF4FFF1FFFF6D8FF809F5A4.taxon	activity	Activity patterns. Like the Diana Monkey, Roloway Monkeys are active, high canopy specialists. They are extraordinarily agile, foraging among terminal branches of emergent canopy trees and making acrobatic leaps to catch insect prey.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF4FFF1FFFF6D8FF809F5A4.taxon	biology_ecology	Movements, Home range and Social organization. Projected similarities between Roloway and Diana Monkeys suggest that groups of Roloway Monkeys contain one adult male, several adult females, and young. Only one social group of the Roloway Monkey has been carefully studied: the group had 14 - 15 individuals (one adult male, six adult females, and young). It traveled 1 - 3 km / day in a home range of 190 ha. The group traveled further per day in the dry season than the wet season. Groups of Roloway Monkeys probably defend territories, although no specific information on territorial behavioris available.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF4FFF1FFFF6D8FF809F5A4.taxon	conservation	Status and Conservation. CITES Appendix I. Classified as Endangered on The IUCN Red List (as C. diana roloway). The Roloway Monkey is listed as Class A in the African Convention on the Conservation of Nature and Natural Resources. It originally occupied forests from the Sassandra River in western Ivory Coast to the Pra River in Ghana, and possibly extending into Togo. It was not found in Togo in recent primate surveys there. The most recent surveys indicate that it is more threatened than has been previously recognized; individuals have been reported at only one site, Tanoé Forest in Ivory Coast. If this is correct, the Roloway Monkey should be recognized as one of Africa’s most endangered primates, and it should accordingly be updated to critically endangered. Unconfirmed reports suggest that it might still hang on in very small numbers at Ankasa in Ghana; a thorough survey of Ankasa should be a high priority. The main threats to the Roloway Monkey include hunting and habitat degradation and loss. The loss of tall mature forest to logging and agriculture has not only degraded preferred habitat but also opened the forest up to commercial hunting for bushmeat. The relatively large size of the Roloway Monkey and the value ofits meat and skin make it a preferred game species. A couple of decades ago, the Roloway Monkey inhabited several parts of West Africa, including Krokosua Hills, Yoyo River, Boin River, and Draw River forest reserves, Ankasa Game Reserve, and Foret Classée de Yaya in Ivory Coast, and Dadieso Forest Reserve and Tanoé Forest in Ghana. Because of high human density around these areas, the Roloway Monkey's habitat has suffered one of the highest recent rates of deforestation in the world. Prior to this, Ghana's 300 km * Bia National Park (the only site where Roloway Monkeys have been studied) was downgraded to a game production reserve. Logging concessions were granted in 1977, and the small remaining park area did not receive adequate protection; no Roloway Monkeys occur there anymore. The Roloway Monkey has been steadily extirpated from both unprotected and protected areas, and it is nearing extinction.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF4FFF2FAF86F8FFB1DF509.taxon	materials_examined	Sudan, White Nile. According to E. Schwarz in 1928, probably Niam Niam or Monbottu country.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF4FFF2FAF86F8FFB1DF509.taxon	discussion	In their 1999 guide to Central African primates, A. Gautier-Hion and colleagues described C. neglectus as belonging to the lhoesti species group. In his 2001 review, C. P. Groves assigned it to its own species group. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF4FFF2FAF86F8FFB1DF509.taxon	distribution	Distribution. S Cameroon (from just N of the Sanaga River, between the Mbam and Djerem rivers), S Central African Republic, Equatorial Guinea, Gabon (S to the Mpassa River and its tributaries), Republic of the Congo (S to the Nambouli River in the Lefini Faunal Reserve), E to DR Congo, Uganda, Kenya in the E of the Rift Valley (Mathews Range Forest Reserve of Samburu), and SW Ethiopia, and S to NE Angola; current presence in Uganda is unknown, but it was, in the 1960 s reported in Bwamba, Bukedi, and Busia districts, and its continued existence in Ethiopia is uncertain.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF4FFF2FAF86F8FFB1DF509.taxon	description	Descriptive notes. Head — body 54 cm (males) and 44 cm (females), tail 70 cm (males) and 59 cm (females); weight 5 - 5 kg (males) and 3.2 - 3.6 kg (females). De Brazza’s Monkey is markedly sexually dimorphic, with females being ¢. 57 % of the weight of males. It is a robust species, with a shorttail. Body is gray with white bands, limbs are dark, arms are black, and there is a white haunch stripe. Underside is dark. It has a black tail. Hairs on the body have a white base, with six or seven alternating bands of white and black. There is a broad orange, crescent-shaped band on its forehead, margined in black. It has a white beard and white fur around its mouth. Scrotum is blue. There are striking age differences in appearance. Newborns are yellow-brown, with yellow limbs and underside. Infants have a dark crown and median dorsal line and a brow band thatis pale yellow. Their cheeks are pale, tail is yellow, and beard is incipient. Infant’s hair is gray at the base, with yellow tips. Juveniles are agouti grayish brown, with a chestnut brow band, rump, and tail base. Occasionally, an individual retains this coloration into adulthood — the “ ezrae ” morph.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF4FFF2FAF86F8FFB1DF509.taxon	biology_ecology	Habitat. Lowland and submontane tropical moist forest, swamp forest, semi-deciduous forest, and Acacia-dominated forest; primarily riparian forests, generally preferring dense vegetation nearrivers. Seasonal flooded forests are also used. De Brazza’s Monkeys sometimes enter monoculture plantations. They have been recorded at elevations of 450 - 1650 m. The highest known elevation recorded is an isolated group in the Mathews Range Forest Reserve of Samburu at 2200 m.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF4FFF2FAF86F8FFB1DF509.taxon	food_feeding	Food and Feeding. De Brazza’s Monkey is mainly frugivorous, with 33 - 74 % of the diet comprised of fruits. Leaves (9 -: 0 - 19 - 5 %) and animal prey — insects (5 % - 10 %) and reptiles — also form a substantial part oftheir diet. They frequently forage on the ground, spending as much as 30 % of their day on the forest floor. They have been reported eating soil.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF4FFF2FAF86F8FFB1DF509.taxon	breeding	Breeding. Sexual maturity is reached at 4 - 6 years. The gestation period is 5 - 6 months, and usually a single infant is born. Only the female looks after the young. Individuals live for up to 22 years. In captivity, infant De Brazza’s Monkeys are reported to mature faster than other guenons.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF4FFF2FAF86F8FFB1DF509.taxon	activity	Activity patterns. De Brazza’s Monkeys are diurnal and semi-terrestrial, spending 50 - 70 % of their time below 5 m high in the forest canopy and ¢. 20 % on the ground. Locomotion is quadrupedal, with associated leaping and climbing.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF4FFF2FAF86F8FFB1DF509.taxon	biology_ecology	Movements, Home range and Social organization. De Brazza’s Monkeys are found in small family groups in Gabon, northern Congo, Bateke Plateau, and central Cameroon. In the 1980 s, it was suspected that they were monogamous, a mating system that would be inconsistent with their sexual dimorphism, which would otherwise signal polygyny. It was subsequently discovered that their mating / social system ranges from facultative monogamy to male-dominant, polygynous, unimale-multifemale groups typical of other guenons. It was supposed that their marked sexual dimorphism reflected a holdover of dimorphism established earlier in their evolutionary history when the degree of polygyny was higher, and that their inclination toward small groups and monogamy was a consequence of selection toward small group size. This is possibly an anti-predator strategy, associated with the fact that they do not form polyspecific associations as do other species of Cercopithecus and are generally more inconspicuous than other forest guenons. One may also suppose that small groups are related to the abundance and dispersion of their food sources and, as such, to avoiding interspecific feeding competition. Solitary males are observed. Group sizes are 2 - 35 individuals. In Trans-Nzoia and West Pokot districts of western Kenya, where they are threatened with extirpation, group sizes are significantly smaller, reported to be only 2 - 6 individuals. Home ranges are 4 - 10 ha, and average daily movements are 500 - 530 m.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF4FFF2FAF86F8FFB1DF509.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Least Concern on The [UCN Red List. De Brazza’s Monkey is listed as Class B in the African Convention on the Conservation of Nature and Natural Resources. Although it is locally threatened by deforestation, fragmentation, hunting, and livestock grazing, it is generally common and widespread. Rates of decline are not believed to be fast enough to warrantlisting in a threatened category.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF7FFF3FF296E2EFCFAFD5C.taxon	materials_examined	Barbary.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF7FFF3FF296E2EFCFAFD5C.taxon	discussion	In his Field Guide to African Mammals published in 1997, J. Kingdon included the following species in his mona superspecies group: C. mona, C. campbell, C. lowe, C. denti, C. wolfi, and C. pogonmias. C. P. Groves in his 2001 Primate Taxonomy followed this same arrangementfor his mona species group. P. Grubb and colleagues in 2003 did likewise, but they considered the form lowe: to be a subspecies of C. campbelli and the form wolfi to be a subspecies of C. pogonias. C. mona hybridizes with C. pogonias; wild hybrids have been observed in Cameroon showing a variety of intermediate coat-pattern features. An albino individual was exhibited at the London Zoo in the early 1960 s. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF7FFF3FF296E2EFCFAFD5C.taxon	description	Descriptive notes. Head-body 41 - 63 cm (males) and 38 - 46 cm (females), tail 64 — 88 cm (males) and 53 - 66 cm (females); weight 4: 4.7 - 5 kg (males) and 2.5 - 4 kg (females). The Mona Monkeyis a heavily built guenon, with a big head and long, thin tail that is normally carried in a high arc toward the back. It has a rich chestnut-colored back and white undersides (extending to inner surfaces of limbs). It can be readily distinguished by two conspicuous white oval patches, one on either side of base of tail. Lower limbs are gray or black on the outside, and callosities are dark gray. Crown of the head is dark gray, speckled with yellow-gold, and there is a white brow band. Cheeks are yellowish and densely furred, and pink skin is exposed on lips and muzzle. Dark skin on upper nose and around eyes, along with dark hair on temples, gives the Mona Monkey a masked appearance. Scrotum is blue. Newborn infants have a very dark coat.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF7FFF3FF296E2EFCFAFD5C.taxon	biology_ecology	Habitat. Primary and secondary lowland rainforest and mangrove, gallery, and riparian forest. The Mona Monkey is an ecological generalist, able to inhabit all but the most severely degraded of habitats; it can live near human habitation, on farmland, plantations and even in gardens. In parts of eastern Nigeria where little forest remains and most forest anthropoid primates are extinct, the Mona Monkeyis often the only forest monkeystill present.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF7FFF3FF296E2EFCFAFD5C.taxon	food_feeding	Food and Feeding. Mona Monkeys are very frugivorous. Seeds, insects, flowers, and nectar are also important components of their diets. At Lama in Benin, individuals spent ¢. 60 % oftheir time feeding on ripe fruits and unripe seeds, especially of Dialium guineense (Fabaceae) and Diospyros mespiliformis (Ebenaceae) in the dry season when fruit is scarce. In the wet season, insects made up ¢. 18 % of their feeding time, and in the dry season, flowers and nectar made up ¢. 37 %.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF7FFF3FF296E2EFCFAFD5C.taxon	breeding	Breeding. Pairs of Mona Monkey call when mating. Copulation calls have two elements — warbles (described as a continuous whining that rises and falls erratically in pitch) and grunts (choppy, low-toned calls that are repeated rapidly). They are given simultaneously, alternately, or singly during a call series. These calls last for c. 8 seconds and can be heard at distances of more than 200 m. Species of Erythrocebus, Allenopithecus, and Miopithecus and the Sun-tailed Monkey (Allochrocebus solatus) give copulation calls, but the Mona Monkey is the only member of the genus Cercopithecus known to do so. Mona Monkeys appear to breed seasonally. In Lama, Benin, copulation calls have been heard in September — March, and newborn infants have been seen in April-September. In Mungo, Cameroon, copulation calls have been heard in September — December, with infants appearing in April-June. Females become sexually receptive about once a month. They do not show genital swelling. A single young is born after a b — 6 month gestation. The baby is carried ventrally, at first supported by its mother’s hand. Sexual maturity is reached at 4 - 6 years. Individuals can live more than 20 years.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF7FFF3FF296E2EFCFAFD5C.taxon	activity	Activity patterns. The Mona Monkeyis diurnal and mostly arboreal. Individuals spend most of their time in the lower layers of the forest canopy, and quite frequently, they move on the ground. In the Southern Bakundu Forest Reserve, Cameroon, 60 % of their day was spent below a height of 15 m. In the Mungo Forest Reserve, Cameroon, Mona Monkeys were most often seen at heights of 4 - 10 m and foraged on the ground more than other species of Cercopithecus. In the Lama Forest, Benin, 29 % of sightings of Mona Monkeys were on the ground. Much of their movement consists of relatively slow and deliberate quadrupedal walking, with infrequent leaping. In all-male groups on Grenada, resting occupied 48 % of the activity budget, feeding and foraging 26 %, grooming 12 %, and moving 8 %. A group was once observed swimming across a 4 mwide creek in a Cameroon mangrove swamp.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF7FFF3FF296E2EFCFAFD5C.taxon	biology_ecology	Movements, Home range and Social organization. Social groups of Mona Monkeys contain 3 - 35 individuals. In Lama, Benin, average group size has been found to be 13 - 5 individuals. Groups include both mixed and all-male groups; no other guenon species forms closely bonded all-male groups. In mixed groups, one or more adult males are accompanied by several adult females and young. Single-sex groups most commonly take the form of a male pair, or a group of males comprising adults, subadults, and juveniles. Up to five males have been seen traveling together in Lama. There is extensive overlap of group ranges in Lama and Mungo forest. Interactions between groups are aggressive. Males produce loud calls and give head-bobbing and branch-shaking displays. Individuals chase each other and squabble, including biting and wrestling. These intergroup encounters are accompanied by a characteristic call, the “ chortle. ” Female Mona Monkeys are most often seen grooming others, whereas adult males are the most frequently the recipients of grooming. Ranging behavior has not been studied in the Mona Monkey.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF7FFF3FF296E2EFCFAFD5C.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Least Concern on The IUCN Red List. The Mona Monkey is listed as Class B in the African Convention on the Conservation of Nature and Natural Resources. Although its habitat is now highly fragmented and degraded across much of its distribution, the Mona Monkey remains relatively widespread and common. It has a remarkable ability to adapt to a wide variety of degraded habitats. Individuals are able to thrive in swamp forests, young secondary forest, and savanna gallery forest. Indeed, this is the most frequently encountered monkey in the heart ofits distribution in southern Nigeria, where other forest monkeys are now rare or absent. The Mona Monkey is therefore not believed to be declining at any rate that would warrant listing in a threatened category. It is occasionally killed as an agricultural pest, but it is especially numerous where hunting pressure is light, such as in Muslim areas in the northern part ofits distribution. The Mona Monkey is present in many protected areas including Fazao-Malfakassa National Park in Togo; Afi Mountain Wildlife Sanctuary in Nigeria; Tafi Atome Monkey Sanctuary in Ghana; Cross River, Gashaka-Gumti, and Okomu national parks in Nigeria; and Mount Cameroon National Park, Takamanda Forest Reserve, Banyang-Mbo Wildlife Sanctuary, and the proposed Ebo National Park in Cameroon.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF6FFF3FFE267F9F9F5FA94.taxon	materials_examined	Sierra Leone.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF6FFF3FFE267F9F9F5FA94.taxon	discussion	In his Field Guide to African Mammals published in 1997, J. Kingdon included the following species in his C. mona superspecies group: C. mona, C. campbell, C. lower, C. denti, C. wolfi, and C. pogonias. C. P. Groves in his 2001 Primate Taxonomy followed this same arrangement for his C. mona species group. P. Grubb and colleagues in 2003 did likewise, but they considered the form lowe: to be a subspecies of C. campbell. Guenons with intermediate features of C. campbelli and C. lowei have been found between the Cavally and Sassandra-Nzo rivers in eastern Ivory Coast. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF6FFF3FFE267F9F9F5FA94.taxon	distribution	Distribution. Senegal (from just N of Casamance River), Gambia, Guinea-Bissau (including Caravela I in the Bijagos Archipelago), Guinea, Sierra Leone, and Liberia (just to Cavally River on the Liberia — Ivory Coast border).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF6FFF3FFE267F9F9F5FA94.taxon	description	Descriptive notes. Head-body 43 - 54 cm (males) and 36 - 43 cm (females), tail 49 - 85 cm (males) and 58 - 68 cm (females); weight 3 - 9 — 4 - 6 kg (males) and c. 2 - 2 kg (females). Campbell's Monkey is similar in appearance to the Mona Monkey (C. mona) and is not immediately identifiable by one particular striking feature ofits coat. Rather, Campbell’s Monkeyis best distinguished from other sympatric forest guenons by lack of white spots on base oftail and its lack of a white nose. Upper back and shoulders are speckled gray-yellow, but lower back, outer surfaces of limbs, and dorsal surface oftail are gray-black. Underside and inner surfaces of limbs are silvery grayish-white. There is a distinct brow band of white hairs, with black tips and a black line running from eye to ear. Cheek whiskers are speckled pale yellow. Face is blue-gray (paler around eyes), and lips and skin of muzzle are pink. Scrotum is blue.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF6FFF3FFE267F9F9F5FA94.taxon	biology_ecology	Habitat. Primary and secondary lowland and riparian forest, mangroves, and woody savanna (e. g. in the Comoé National Park in northern Ivory Coast). Campbell's Monkey can also be found on subsistence agricultural land and in young secondary growth, bush, and scrubby vegetation referred to as “ farmbush. ”	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF6FFF3FFE267F9F9F5FA94.taxon	food_feeding	Food and Feeding. Fruit is the main constituent of the diets of Campbell's Monkeys, and they also eat leaves, flowers, insects, and other small animals. In Tai Forest National Park, Ivory Coast, individuals spend 46 % of their feeding time eating fruits, especially those of Sacoglottis gabonensis (Humiriacae), Dialium aubrevillei (Fabaceae), and Memecylon lateriflorum (Melastomataceae). About 33 % of feeding time is dedicated to foraging for insects. Campbell's Monkeys carefully search through foliage for rolled up leaves and in rotting wood for insect larvae. They have been seen to eat spider webs. They also eat leaves; 8 % of feeding time can be taken up eating foliage. When in mangroves, they eat leaves, seeds, crabs, shrimps, and mudskippers. They are a serious farm pest in some areas.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF6FFF3FFE267F9F9F5FA94.taxon	breeding	Breeding. Ovulatory cycles of Campbell's Monkeys last about a month. They do not exhibit sexual swelling. In Ivory Coast, mating takes place in June-September, and births occur in November — January at the end of the long rainy season and in the early dry season. The gestation period is c. 180 days. A single young is born each year. The infant is carried ventrally, at first helped by the mother. Weaning takes place when offspring are c. l year old, and sexual maturity is reached at 4 - 6 years. Instances of infanticide have been observed.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF6FFF3FFE267F9F9F5FA94.taxon	activity	Activity patterns. Campbell's Monkey is diurnal and predominantly arboreal. In the Tai Forest, individuals spend 70 % of their time feeding and foraging, 20 % resting, and 7 % traveling. Campbell’s Monkeys spend more time in the lower parts of the forest canopy and walk more and leap less than any other arboreal guenons. Further studies in the Tai Forest have shown that they use the ground for 15 % of their time, and the shrub layer for 22 % oftheir time. About 58 % oftheir feeding time is spent in the shrub layer (below 5 m) and on the ground. Campbell’s Monkeys also sit when resting and stand when feeding, more than any other monkey. They are reported to be good swimmers, allowing them to reach small islands to forage.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF6FFF3FFE267F9F9F5FA94.taxon	biology_ecology	Movements, Home range and Social organization. Average group sizes of Campbell’s Monkeys are 9 - 14 individuals, although groups of up to 33 have been reported. Typically, groups contain a single adult male and several adult females. Males that are not integrated into heterosexual groups form all-male groups that include adults and subadults. Aggressive male takeovers of one-male groups have been reported. They are accompanied by infanticide. Campbell's Monkey show a relatively high level of affiliative interaction among individuals in groups, manifested mostly by grooming between adult females. Low levels of agonistic behavior within groups are attributed to small group sizes and their dispersed feeding patterns. Interactions between groups, although infrequent, are often aggressive. During such encounters, males show more aggressive behavior (threats and loud calling) than females. In a day, groups will travel 1000 - 1200 m. In Tai Forest, annual home range sizes are 40 - 80 ha. Predators include crowned hawk-eagles (Stephanoaetus coronatus), Leopards (Panthera pardus), and Chimpanzees (Pan troglodytes). Campbell's Monkeys give alarm calls that are specific to Leopards and a different one for eagles. They also make loud calls when they are alarmed, such as when they hear a tree fall, but in these cases, calls are introduced by two boom calls that evidently signal that loud calls do not concern presence of a predator. Males have short bouts of loud calling (also, introduced by two boom calls) every evening, at dusk, which have to do with defense of their group and their territory.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF6FFF3FFE267F9F9F5FA94.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Least Concern on The [UCN Red List. Campbell's Monkey islisted as Class B in the African Convention on the Conservation of Nature and Natural Resources. Despite the fact that Campbell's Monkey has been affected by habitat loss through deforestation, it can thrive in disturbed habitats and remains common in many parts ofits distribution. It has a large distribution and an ability to adapt to a wide variety of habitats. It is hunted but, along with the Spot-nosed Monkey (C. petaurista), it still remains one of the most abundant primates in the Upper Guinea forest zone. Campbell’s Monkey occurs in most protected areas in its distribution, including Abuku Nature Reserve in The Gambia, Tai Forest National Park in Ivory Coast, Sapo National Park in Liberia, Basse Casamance National Park in Senegal, and Tiwai Island Wildlife Sanctuary in Sierra Leone.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF6FFFCFAEC60A1FBAAFB81.taxon	materials_examined	Ivory Coast, Bandama.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF6FFFCFAEC60A1FBAAFB81.taxon	discussion	In his Field Guide to African Mammals published in 1997, J. Kingdon included the following species in his C. mona superspecies group: C. mona, C. campbelli, C. lowes, C. denti, C. wolf, and C. pogonias. C. P. Groves in his 2001 Primate Taxonomy followed this same arrangement for his mona species group. P. Grubb and colleagues in 2003 did likewise, but they considered the form lowei to be a subspecies of C. campbelli. Groves classified lowei as a distinct species, which is followed here. C. campbelli x C. lowei intermediates occur where the species replace each other in an area between the Cavally and Sassandra riversin Ivory Coast. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF6FFFCFAEC60A1FBAAFB81.taxon	distribution	Distribution. Patchily from the Cavally River in W Ivory Coast, E to the Volta River in Ghana.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF6FFFCFAEC60A1FBAAFB81.taxon	description	Descriptive notes. Head-body 45 - 52 cm (males) and 40 - 43 cm (females), tail 66 — 75 cm (males) and 54 - 59 cm (females); weight 5.2 - 5.8 kg (males) and 2.3 - 8 kg (females). Male Lowe’s Monkeys are much larger than females. Crown is chestnut flecked with yellow. It has a yellow brow band with black-tipped hairs. Cheek whiskers are speckled yellow, with gray bases. Ear tufts are grizzled and often yellowish. Face is bluegray exceptfor lips and skin of the muzzle, which are pink. Dorsum is grizzled brown, slightly orange or golden-red with a contrastingly dark gray to blackish rump. Underparts are white. Outer thighs are dark gray and forearms to above elbows are blackish. Base and distal one-third oftail are black, but otherwise tail is pale yellowish.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF6FFFCFAEC60A1FBAAFB81.taxon	biology_ecology	Habitat. A wide range of lowland habitats, including primary evergreen and semideciduous forest, secondary growth forest, gallery forest, and forest-savanna mosaics north of the rainforest zone. Lowe’s Monkeys are uncommon in marshy areas and mangroves but can sometimes be seen in secondary “ farmbush ” (secondary growth and scrub on farmland). They are most often observed in the middle and lower levels of the forest canopy, rarely using emergents. The average height above the ground for foraging groups is c. 7 m. Lowe’s Monkeyis a lowland species; it can be found at elevations up to 300 m, but the majority ofits distribution is below 150 m.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF6FFFCFAEC60A1FBAAFB81.taxon	food_feeding	Food and Feeding. Diets of Lowe’s Monkeys are mainly fruit, including figs (Ficus, Moraceae) and fruits of Cola (Sterculiaceae) and Baphia (Fabaceae). They also eat leaves, seeds, flowers, bark, and insects (larvae and orthopterans). In one area, arthropods dominated the diet, 56 - 63 %, with cultivated fruits making up 24 - 26 %. Lowe's Monkeys drink water by dipping their hands into tree holes and licking theirfur.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF6FFFCFAEC60A1FBAAFB81.taxon	breeding	Breeding. Female Lowe’s Monkeys reach sexual maturity at c. 3 years of age and males at c. 4 years. The gestation period is 147 - 180 days. Weaning occurs at c. 18 months. Seasonal mating has been observed in central southern Ivory Coast in June-December, and births occur in November — January to coincide with the growing season. Both allomothering and exclusive maternal care have been observed. Infanticide has been recorded. Life span is ¢. 28 years.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF6FFFCFAEC60A1FBAAFB81.taxon	activity	Activity patterns. Lowe’s Monkeys are diurnal and arboreal. Locomotion is primarily quadrupedal, with climbing and leaping, and bipedal postures have also been recorded, especially when foraging for insects.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF6FFFCFAEC60A1FBAAFB81.taxon	biology_ecology	Movements, Home range and Social organization. Lowe's Monkeys live in unimalemultifemale groups, with 4 - 5 females and their offspring; generally, there are 6 - 12 individuals in a group, averaging c. 10 individuals. Solitary males have also been observed. Males and females disperse from their natal groups. A group may split into subgroups when foraging. Average home rangeis c. 40 ha, but a group using a remnant forest had smaller ranges of 3 - 14 ha. Lowe’s Monkeys are often associated with other species, including the Diana Monkey (C. diana). Lowe’s Monkeysinitiate and terminate such associations, presumably gaining anti-predator benefits. Predators include crowned hawk-eagles (Stephanoaetus coronatus), Leopards (Panthera pardus), and Chimpanzees (Pan troglodytes).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF6FFFCFAEC60A1FBAAFB81.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Least Concern on The [UCN Red List (as C. campbelli lower). Lowe’s Monkey is listed as Class B in the African Convention on the Conservation of Nature and Natural Resources. It is widespread and adaptable when its forests are fragmented and degraded, as long there is some cover and connectivity among forest patches. Lowe’s Monkey is heavily hunted and reported to be declining as a result, but declines are not thought to be at a pace that would warrant listing in a threatened category. It has been extirpated in Banco National Park, Ivory Coast. It survives well in sacred forests in both Ghana and Ivory Coast and occurs in Kakum National Park and Boabeng-Fiema Monkey Sanctuary in Ghana.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF9FFFCFF2161D6F851F75E.taxon	materials_examined	Equatorial Guinea, Bioko Island.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF9FFFCFF2161D6F851F75E.taxon	discussion	In his Field Guide to African Mammals published in 1997, J. Kingdon included the following species in his C. mona superspecies group: C. mona, C. campbelli, C. lowe, C. denti, C. wolfi, and C. pogonias. C. P. Groves in his 2001 Primate Taxonomy followed this same arrangement for his mona species group. P. Grubb and colleagues in 2003 did likewise, but they considered the form wolfi to be a subspecies of C. pogonias. A fourth subspecies schwarzianus named by H. Schouteden in 1946 from Mayumbe in DR Congo, similar to gray: but darker, is recognized by some authors. Three subspecies recognized.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF9FFFCFF2161D6F851F75E.taxon	distribution	Subspecies and Distribution. C. p. pogoniasBennett, 1833 — SENigeria (fromtheCrossRiver), NWCameroon (NofSanagaRiver), andBiokoI (EquatorialGuinea). C. p. grayiFraser, 1850 — SCameroon (SofthemiddleSanagaRiver), SWCentralAfricanRepublic, NWDRCongo (EoftheUbangiRiver, WtotheItimbiriRiverBasinNoftheCongoRiver), NEGabon, NRepublicoftheCongo, andpossiblyAngola (Cabinda). C. p. nigripes Du Chaillu, 1860 — Equatorial Guinea, W Gabon, Republic of the Congo (between the Ogooué and Congo rivers).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF9FFFCFF2161D6F851F75E.taxon	description	Descriptive notes. Head — body 50 - 57 cm (males) and 44 - 48 cm (females), tail 69 - 87 cm (males) and 60 - 74 cm (females); weight 4 - 5 — 4 - 7 kg (males) and 3.1 - 3.5 kg (females). As the common name suggests, the Crowned Monkey is distinguished by a crest of black hair on the crown of the head. The “ Mohawk ™ shaped crest runs medially, dividing patches of speckled yellow fur on either side of the crown. Dark color on upper nose and around eyes, along with dark hair on temples, also gives the Crowned Monkey a “ masked ” appearance. Ear tufts are prominent and pointed. Lower muzzle is flesh-colored, and ear tufts are orange or yellow. Back is variably agouti-gray, and a dorsal band of darker hair runs from shoulders to base oftail, broadening posteriorly. In contrast, underside, including chest, belly, and inner limb surfaces, are bright golden-orange. Outer surfaces of limbs are variably speckled gray or black. Tail is dark above and golden-brown below for about two-thirds ofits length, becoming entirely black at the tip. The “ Golden-bellied Crowned Monkey ” (C. p. pogonias) has a sharply defined black saddle. Its flanks are speckled with white, and its lower limbs are yellowish, with only toes being black. Median black-striped crest on the crown extends forward to brows. “ Gray’s Crowned Monkey ” (C. p. grayi) can be distinguished by marginally different coloration: dark chestnut-red above with a dark-red saddle and a lemon-yellow or whitish underside. Its flanks and thighs are orange, and its lower limb segments are buffy-gray with only toes black. Cheeks and ear tufts are lemon-yellow or whitish. As with the Golden-bellied Crowned Monkey, its crest extends forward to brows. The “ Black-footed Crowned Monkey ” (C. p. nigripes) is similar to the Goldenbellied Crowned Monkey in having a dark-gray upper body, with a sharply defined black saddle and an orange or yellow underside, but with lower limb segments buffygray, whitish speckling on flanks, and entirely black hands and feet. Its crest ends well above brows, so that yellow areas meet in the midline and form a U-shape.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF9FFFCFF2161D6F851F75E.taxon	biology_ecology	Habitat. Lowland tropical forest, especially primary high-canopy forest, but also mature secondary forest up to submontane and montane elevations, and gallery forest extending into savannas. In Idenau, Cameroon, the Crowned Monkey uses higher canopy levels than does any other species. About 50 % of observations of the Crown Monkeys in Idenau were at heights of 17 m or more above the ground. In southeastern Bioko, they were most frequently seen at heights of 10 - 24 m. On the edge of Mount Cameroon, they were most abundant in mature forest, moderately abundant in older secondary forest, and absent in young secondary forest.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF9FFFCFF2161D6F851F75E.taxon	food_feeding	Food and Feeding. The Crowned Monkey feeds mainly on fruits and seeds, supplemented with leaves, shoots, flowers, and insects (especially orthopterans). In Lopé National Park, Gabon, the Black-footed Crowned Monkey spent 78 % ofits feeding time on fruits and seeds, 9 % on flowers, 6: 5 % on leaves, and 6 - 5 % on animal matter. In Makokou, Gabon, stomach contents of Gray’s Crowned Monkeys killed by hunters contained, on average, 78 % (by dry weight) fruits and seeds, 12 % animal remains (beetles, caterpillars, and grasshoppers), and 6 % leaves. The Golden-bellied Crowned Monkey feeds in higher levels in the forest canopy than other guenon species. In Southern Bakundu Forest Reserve, Cameroon, 50 % of feeding heights were above 21 m. Despite its propensity to use the upper canopy, it goes to the ground to raid Crops in some areas.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF9FFFCFF2161D6F851F75E.taxon	breeding	Breeding. There is no specific information for this species, but Gray’s Crowned Monkey in north-eastern Gabon gives birth in November — April, coinciding with the driest time of the year.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF9FFFCFF2161D6F851F75E.taxon	activity	Activity patterns. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF9FFFCFF2161D6F851F75E.taxon	biology_ecology	Movements, Home range and Social organization. Social groups of Crowned Monkeys are 9 - 19 individuals (average 13 - 14) in south-western Cameroon. They are most commonly composed of a single adult male and several adult females. Observations indicate that Crowned Monkeys move over wide areas. In Idenau, Cameroon, groups of Crowned Monkeys move over areas two to three times as large as groups of Puttynosed Monkeys (C. nictitans) in the same forest. Movements are influenced by fruit abundance and are therefore seasonal. In Central African forests, groups of Crowned Monkeys have home ranges of at least 100 ha, with travel distances of 1500 m / day. The adult male has a characteristic booming call, uttered when the group is dispersed. They also give territorial calls referred to as “ hacks. ”	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF9FFFCFF2161D6F851F75E.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Least Concern on The [UCN Red List, including the subspecies gray: and nigripes, but the nominate subspecies pogonias is classified as Vulnerable. The Crowned Monkeyis listed as Class B in the African Convention on the Conservation of Nature and Natural Resources. It has been affected in parts of its distribution by habitat loss through deforestation, agriculture, and human settlement, and it is also hunted for meat. It is relatively widespread and is present in several protected areas, including Korup National Park in Cameroon (an important stronghold to ensure its survival in the Cameroon-Nigeria border area), Odzala National Park in Republic of the Congo, Monte Alen National Park in Equatorial Guinea, and Salonga National Park in DR Congo. Although the Crowned Monkey is one of the smaller species of Cercopithecus, it might be more vulnerable to hunting than other species because ofits relatively narrow feeding and habitat requirements. Nevertheless, based on its widespread range and slow rate of decline, the Crowned Monkey is not thought to be threatened. Although no major declines have been reported on Bioko Island, the Golden-bellied Crowned Monkey on the mainland is heavily hunted in many parts of its distribution where anti-poaching patrols are absent. This heavy hunting, combined with habitatloss, has lead to a decline of ¢. 30 % over the past 27 years (three generations).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF9FFFDFA2F6DF6FC79F66E.taxon	materials_examined	Central West Africa.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF9FFFDFA2F6DF6FC79F66E.taxon	discussion	In his Field Guide to African Mammals published in 1997, J. Kingdon included the following species in his C. mona superspecies group: C. mona, C. campbell, C. lowe, C. dents, C. wolf, and C. pogonias. C. wolfi is considered as a subspecies of C. pogonias by some authorities, including P. Grubb and colleagues in their 2003 review. C. P. Groves considered it to be a distinct species, with three subspecies: wolfi, pyrogaster, and elegans. A. Gautier-Hion and colleagues in their guide to Central African primates published in 1999 included denti as a fourth subspecies. Here, Grovesis followed by recognizing wolfi as a distinct species. Three subspecies recognized.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF9FFFDFA2F6DF6FC79F66E.taxon	distribution	Subspecies and Distribution. C. w. wolfiMeyer, 1891 — DRCongo, betweentheCongoandSankururivers; StotheKasaiRiver (7 ° 30 ” S); theCongo-LualabariversystemappearstobethemajorbarrierintheNandE. C. w. elegans — Dubois & Matschie, 1912 — DRCongobetweentheLomamiandLualabarivers (2 — 4 ° S), StothesavannaandNtothebarrierofaswampforestthatseparatesitfromthesubspecieswolfi. C. w. pyrogaster Lonnberg, 1919 — DR Congo S of Kasai River, limited to a narrow area between the Kwango River in the W and the Lulua River, a tributary of the Kasai River, in the E.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF9FFFDFA2F6DF6FC79F66E.taxon	description	Descriptive notes. Head — body 28 - 54 cm (males) and 46 cm (females), tail 71 - 72 cm (males) and 60 - 66 cm (females); weight 3.8 - 6.5 kg (males) and 2.8 - 3.7 kg (females). Males are larger than females. Wolf’s Monkey is dark gray and generally redder on the back. Arms are black, legs are reddish, and underside is sharply demarcated white, yellow, or red. Terminal one-third to half of the tail is black. Wolfs Monkey has a pale brow band, two lateral black crown stripes, and red or white ear tufts. Body hairs are gray at the base, with three or four alternating light / dark bands. Scrotum is blue. The “ Congo Basin Wolf’s Monkey ” (C. w. wolfi) has a medium dorsal chestnut patch, red legs, a yellow underside sometimes with an orange stripe along flanks, yellow cheek whiskers speckled with black, and red ear tufts. The “ Fire-bellied Wolf's Monkey ” (C. w. pyrogaster) is similar to the Congo Basin Wolf’s Monkey but more deeply colored. Its arms are speckled with yellowish white, its underside is red, and its cheek whiskers are buffy, speckled with black. In the “ Lomami River Wolfs Monkey ” (C. w. elegans), back is brown becoming darker toward rump, forearms are black with pale speckling on upper arms, legs are light gray, underside is white, cheek whiskers are yellowish-white with dark speckling, darker posteriorly, and ear tufts are white.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF9FFFDFA2F6DF6FC79F66E.taxon	biology_ecology	Habitat. The Congo Basin Wolf's Monkey lives in a variety of forest types, including high-canopy tropical evergreen primary forest and secondary, seasonal, and swamp forest. The scant literature available suggests that it occurs at 350 - 500 m above sea level. It is also reported in Mbau forest (Gilbertiodendron deweuvrei, Fabaceae, monodominant forest) in Lomako, Caesalpinoideae-dominated forest in Salonga National Park, seasonally flooded forest, and slope forest. The Lomami River Wolf’s Monkey occurs in riparian forest and savanna mosaics. Habitats of the Fire-bellied Wolf’s Monkey have not been described in the literature.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF9FFFDFA2F6DF6FC79F66E.taxon	food_feeding	Food and Feeding. Several studies of feeding behavior of the Congo Basin Wolf's Monkey report a diet offruit (27 - 47 %), seeds (20 - 49 - 8 %), young and mature leaves (20 - 29 %), flowers (5 — = 11 %), and insects (20 %). The diets of Lomami River and Firebellied Wolf's monkeys have not been explicitly described, but they are probably similar to that of the Congo Basin Wolf's Monkey.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF9FFFDFA2F6DF6FC79F66E.taxon	breeding	Breeding. There is no specific information available for this species. The gestation period of a captive individual was 170 days.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF9FFFDFA2F6DF6FC79F66E.taxon	activity	Activity patterns. Wolf's Monkeys are diurnal and highly arboreal. In the Lomako Forest, DR Congo, they usually travel and forage in the middle and upper forest canopy at heights of c. 17 m, and they have never been seen below 5 m. They spend ¢. 65 % of their time at heights of 5 - 20 m and 35 % at 20 - 40 m. Aspects of locomotion and posture have not been reported, but they are probably similar to the closely related Crowned Monkey (C. pogonias).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF9FFFDFA2F6DF6FC79F66E.taxon	biology_ecology	Movements, Home range and Social organization. Wolf's Monkeys live in hierarchical, male-dominated, unimale-multifemale groups. Groups of 10 - 30 individuals have been recorded in Salonga National Park and Lomako and Malabi forests. Solitary males have also been reported. There is no information available for Lomami River Wolf’s and the Fire-bellied Wolf’s monkeys.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF9FFFDFA2F6DF6FC79F66E.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Least Concern on The [UCN Red List (as C. pogonias wolfi), but the subspecies elegans (as C. pogonias elegans) is classified as Near Threatened. The subspecies pyrogaster has not been assessed. Wolf’s Monkey is listed as Class B in the African Convention on the Conservation of Nature and Natural Resources. The Congo Basin Wolf’s Monkey has a widespread distribution. Although its population is declining, it is notlikely to be declining fast enough to be considered threatened. The Lomami River Wolf’s Monkey is subject to hunting and habitat loss in its very restricted distribution. It is believed that the population has declined 20 - 25 % over the past 27 years, almost qualifying it for a threatened category.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF8FFFDFAF56430F98FF668.taxon	materials_examined	DR Congo, between Mawambi and Avakubi, Ituri Forest.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF8FFFDFAF56430F98FF668.taxon	discussion	In his Field Guide to African Mammals published in 1997, J. Kingdon included the following species in his C. mona superspecies group: C. mona, C. campbelli, C. lower, C. denti, C. wolfi, and C. pogonias. C. dent is regarded as a subspecies of C. pogonias by some authorities, including P. Grubb and colleagues in their 2003 review. The [UCN also lists this taxon as one of six subspecies of C. pogonias, along with the nominate pogonias, nigripes, grayi, wolfi, and elegans. J. - M. Lernould in 1988 and A. Gautier-Hion and colleagues in 1999 classified dent: as a subspecies of C. wolfi. C. P. Groves in his 2001 Primate Taxonomy considered it to be a distinct species, the classification followed here. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF8FFFDFAF56430F98FF668.taxon	distribution	Distribution. E DR Congo (E and N of the Congo-Lualaba river system, extending N to the Itimbiri River and S to the Kasongo savanna), W Uganda, and NW Rwanda.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF8FFFDFAF56430F98FF668.taxon	description	Descriptive notes. Head — body 40 - 50 cm, tail 68 - 90 cm; weight 4.3 - 5.7 kg (males) and 2.8 - 3.4 kg (females). Male Dent’s Monkeys are larger than females. Back is a rich grizzled mahogany brown. Hairs are gray at the base, with three or four alternating light / dark bands. Head is yellowish. Forelimbs are blackish, and legs are a dark grizzled gray. Underside and inner side of limbs are a sharply contrasting white. Skin offace is bluegray except around the muzzle, which is pale. Dent’s Monkey has a pale brow band and black lateral crown stripes. Tail is gray with a black tip.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF8FFFDFAF56430F98FF668.taxon	biology_ecology	Habitat. In a study carried out in the Ituri Forest in 1991, Dent's Monkey was occasionally present in Mbau and riparian forests but more frequently in mixed forest. It showed a preference for secondary forest because of the year-round abundance of fruit. Primary forest is used during short seasonal fruiting peaks. They use the middle forest canopy most frequently. They are also reported in swamp forest in the Ituri at elevations of 700 - 1000 m.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF8FFFDFAF56430F98FF668.taxon	food_feeding	Food and Feeding. Dent’s Monkeys are primarily frugivore-insectivores. They also eat leaves. In Bwamba, Uganda, they feed on shoots.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF8FFFDFAF56430F98FF668.taxon	breeding	Breeding. There is no specific information available for this species. One captive individual lived 14 years.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF8FFFDFAF56430F98FF668.taxon	activity	Activity patterns. Dent's Monkeys are diurnal and highly arboreal. They are canopy dwellers, spending c. 30 % of their time at heights of 20 - 40 m or higher and ¢. 20 % at 5 - 20 m. Locomotion is quadrupedal.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF8FFFDFAF56430F98FF668.taxon	biology_ecology	Movements, Home range and Social organization. Group sizes of Dent’s Monkeys are 7 - 21 individuals. Their social organization has not been studied in detail, but they are presumably similar to the closely related Crowned Monkey (C. pogonias), having a unimale-multifemale hierarchical social system.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFF8FFFDFAF56430F98FF668.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Least Concern on The [UCN Red List (as C. pogonias denti). Dent's Monkey has a large range, and although its population is declining, it is unlikely to be doing so fast enough to warrant listing as threatened. Nevertheless, because Dent’s Monkey is currently subsumed under the Crowned Monkey by IUCN, it has not been evaluated as a full species, and therefore its threat status is not definitive.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFDFFF8FF3C641AFC6DF30C.taxon	materials_examined	Democratic Republic of the Congo, Ituri Forest.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFDFFF8FF3C641AFC6DF30C.taxon	discussion	C. hamlyni is placed in its own species group. Lowland and highland forms of C. hamlyni differ in cranial morphology and are classified as subspecies: hamlyni and kahuziensis. In their 2003 review of the taxonomy of the African primates, P. Grubb and colleagues acknowledged doubts expressed by J. Hart concerning the validity of kahuziensis and indicated the need for further study. Two subspecies recognized.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFDFFF8FF3C641AFC6DF30C.taxon	distribution	Subspecies and Distribution. C. h. hamlyniPocock, 1907 — EDRCongo, fromthelowlandsoftheRiverLualabaEtoIturiForestandtheAlbertineRiftinWRwanda (NyungweForestNationalPark), andextendingNtotheLindi-Nepokosystem, andStothelimitoftherainforest (poorlydefined); possiblyfoundinSUgandaandNBurundi. C. h. kahuziensis Colyn & Rahm, 1987 — E DR Congo, Mt. Kahuzi in the Albertine Rift Mts.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFDFFF8FF3C641AFC6DF30C.taxon	description	Descriptive notes. Head-body 55 cm, tail 57 cm (males; no measurements for females); weight 5 - 5 kg (males) and 3.1 - 3.7 kg (females). Males are larger than females. The Owl-faced Monkey has dark and dense grizzled gray pelage that is elongated to form a hood over its crown and cheeks. Hairs are gray at the base, with four or five alternating light / dark band pairs. Scrotum is blue. The “ Nose-stripe Owl-faced Monkey ” (C. h. hamlyni) has a narrow white stripe running down its nose from the interocular region to the border of the upper lip, which with a white brow band forms the characteristic “ T ” of white on its face. Its body is olive-green to yellow-gray, varying in tone. There are silver-gray hairs on backs of thighs, perianal region, and base oftail. Its underside, limbs, and distal half of tail are black. The “ Mount Kahuzi Owl-faced Monkey ” (C. h. kahuziensis) has a darker body, an olive-green back, and entirely black facial hairs, except around eyes where they are a lighter grayish. It has no white nasal or brow band. Its tail is shorter than its head-body length.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFDFFF8FF3C641AFC6DF30C.taxon	biology_ecology	Habitat. The Nose-stripe Owl-faced Monkeylives in a variety of forest habitats including lowland and mid-montane moist forest and secondary growth forest. It has been reported in open canopy forest with a dense understory. The Mount Kahuzi Owl-faced Monkey occurs in montane bamboo forest. Owl-faced Monkeys occur at elevations of 600 - 3300 m; the Mount Kahuzi Owl-faced Monkey occurs above 3000 m in both Tshiberimu and Kahuzi.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFDFFF8FF3C641AFC6DF30C.taxon	food_feeding	Food and Feeding. Owl-faced Monkeys eatfruit, seeds, young and mature leaves, flowers, bark, lichen, and insects. In higher elevation habitats, Mount Kahuzi Owl-faced Monkeys eat leaves and shoots of bamboo and leaves, shoots, pith, and stems of wild celery (Peucedanum, Apiaceae) and Vernonia (Asteraceae). They also eat fruits such as blackberries (Rubus, Rosaceae), and those of Podocarpus (Podocarpaceae), Myrica (Myricaceae), Schefflerea (Araliaceae), Dombeya (Malvaceae), and Cassipourea (Rhizophoraceae).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFDFFF8FF3C641AFC6DF30C.taxon	breeding	Breeding. In captivity, neonatal Owl-faced Monkeys weigh 280 g. The time from birth to weaning is ¢. 8 - 5 months. The infant first begins to climb at two weeks. Only the mother cares for the young. There is an apparent birth season in June-October in at least part of its distribution. Life span in captivity is twelve years.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFDFFF8FF3C641AFC6DF30C.taxon	activity	Activity patterns. Owl-faced Monkeys are diurnal and semi-terrestrial. They spend more than 60 % of their time on the ground and show a preference for lower levels of the forest canopy at heights of less than 10 m. Locomotion is quadrupedal. Climbing was reported for 30 % of the time, with vertical clinging and leaping reported at 10 %.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFDFFF8FF3C641AFC6DF30C.taxon	biology_ecology	Movements, Home range and Social organization. Owl-faced Monkeys live in maledominated, unimale-multifemale or multimale-multifemale social groups. Monogamy is facultative. Group sizes are 2 - 22 individuals. Solitary individuals are frequently observed.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFDFFF8FF3C641AFC6DF30C.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Vulnerable on The IUCN Red List, but the two subspecies have not been assessed. The Owl-faced Monkey is listed as Class B in the African Convention on the Conservation of Nature and Natural Resources. It has undergone declines in the past and continues to fall in numbers. Bamboo is often harvested in the montane forests where it occurs. It is rare in DR Congo and Rwanda but relatively abundant in Mount Kahuzi-Biéga National Park. Its entire distribution is in a region of intense conflict, where civil war has exacerbated identified threats. Declines will probably continue in the face of the ongoing political climate of the region, resulting in an overall population reduction of more than 30 % over 30 years (three generations).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFDFFF8FA38641EF90FF558.taxon	materials_examined	West Bank of Lomami River (S 1 - 02237 ° to S 1 - 4280 °, E 24 - 42368 ° to E 25 - 03843 °), Democratic Republic of the Congo.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFDFFF8FA38641EF90FF558.taxon	discussion	Morphological and molecular data show that C. lomamiensis is closely related to, but distinct from, C. hamlyni. Aspects that are shared with C. hamlyni and distinguish them from other guenons include a distinctive skull shape, a prominent facial mane on the cheeks and crown of both sexes, a white-to-cream nose stripe, and boomcall choruses given in the early morning. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFDFFF8FA38641EF90FF558.taxon	distribution	Distribution. EC Congo Basin in DR Congo, from the upper Tshuapa River (24 ° E) in the Sankuru District of Kasai Oriental Province to the Lomami River (25 ° E) in Maniema and Orientale provinces in the E; the N limits are unclear, but it apparently does not occur N of 1 ° S; in the S, its range extends the limits of the forest at c. 3 ° 30 ° S.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFDFFF8FA38641EF90FF558.taxon	description	Descriptive notes. Head — body 47 - 65 cm (males, n = 2), tail 65 - 78 cm (males, n = 2); weight 4.7 - 1 kg (males); no measurements available for adult females. Diagnostic features of the Lesula are a mane of long grizzled blond hairs, framing a pale, naked, pinkish face and muzzle, with a distinct white-to-creamy nose stripe. Hairs over back are banded buff or amber with black. There is a prominent rufous-to-amber median stripe on distal half to one-third of back that extends onto base of tail. Tail is otherwise black. Anterior two-thirds of dorsum, including head and mane, is buff. Chin, throat, and upper ventrum are yellowish buff, contrasting with black lower ventrum and abdomen. Fur on shoulders and forelimbs is black. Face, circumocular region, and eyelids are pinkish. There is a buff diadem. Upper half of thighsis silvery-gray, the remainder is black. Hands and feet are black. Adult male Lesulas have a large, bright, pale-blue patch of skin on their perineum and buttocks. Scrotum is pale blue.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFDFFF8FA38641EF90FF558.taxon	biology_ecology	Habitat. Mature terra firma evergreen humid forests, including mixed forests and forests dominated by Gilbertiodendron dewevrei (Fabaceae), at elevations of 400 - 715 m. The Lesula is sometimes found in regenerating forest around settlements. It does not enter seasonally inundated forest or gallery forest in savannas in the southern part of its distribution.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFDFFF8FA38641EF90FF558.taxon	food_feeding	Food and Feeding. The Lesula eats leaf petioles, ripe and unripe fruits, flower buds, and meristems of two species of terrestrial herbaceous Marantaceae.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFDFFF8FA38641EF90FF558.taxon	breeding	Breeding. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFDFFF8FA38641EF90FF558.taxon	activity	Activity patterns. There is no specific information available for this species, but the Lesula, like the Owl-faced Monkey (C. hamlyni), is semi-terrestrial.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFDFFF8FA38641EF90FF558.taxon	biology_ecology	Movements, Home range and Social organization. The Lesula is not uncommon but is difficult to see, It was detected by researchers mainly by boom calls given as a dawn chorus, as is the habit of the closely related Owl-faced Monkey. Lesulas rarely produce these calls during the day, but sometimes at night. When seen, groups number 1 - 5 individuals. In eleven of 19 encounters, they were with groups of other primates: Wolf's Monkeys (C. wolfi), Red-tailed Monkeys (C. ascanius katangae), or Tshuapa Red Colobus (Piliocolobus tholloni). A little more than one-third of the individuals seen were on the ground, on some occasions scavenging food items dropped by monkeys foraging above them. In one observed encounter a crowned hawk-eagle (Stephanoaetus coronatus) attacked and fatally wound a subadult female Lesula.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFDFFF8FA38641EF90FF558.taxon	conservation	Status and Conservation. CITES Appendix II. The Lesula has not been assessed on The IUCN Red List. The team that discovered and described the Lesula carried out a preliminary assessment and indicated that it should be classified as Vulnerable. The region where it occurs is restricted, and the extent of occurrence is believed to be c. 17,000 km?. This area is remote, and forests remain largely intact. Hunting for subsistence and the bushmeat market (Kindu and Kisangani) since the turn of the century is the main threat. The Lesula is protected in Réserve Naturelle de Sankuru, which along with a proposed national park would cover a large part ofits natural distribution.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFDFFF9FA006FFEFE66F358.taxon	materials_examined	Guinea.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFDFFF9FA006FFEFE66F358.taxon	discussion	C. petauristais a member of the C. cephus superspecies group, as defined by J. Kingdon in his 1997 Field Guide to African Mammals. This group also includes C. cephus, C. sclateri, C. erythrotis, C. ascanius, and C. erythrogaster. C. P. Groves in his 2001 Primate Taxonomy gave the same composition for his cephus species group. Two extant subspecies are divided from one another by the Sassandra River (Ivory Coast), with some hybridization / intergradation reported on the right bank of the Nzo River. Two subspecies recognized.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFDFFF9FA006FFEFE66F358.taxon	distribution	Subspecies and Distribution. C. p. petauristaSchreber, 1774 — C & EIvoryCoast (EoftheSassandraRiver), Ghana, andWTogo; possiblyinBenin. C. p. buettikofer: Jentink, 1886 — SE Senegal, S Guinea-Bissau, S Guinea, Sierra Leone, Liberia, and W Ivory Coast (W of the Nzo-Sassandra River system).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFDFFF9FA006FFEFE66F358.taxon	description	Descriptive notes. Head-body 44 - 53 cm (males) and 40 - 44 cm (females), tail 57 — 79 cm (males) and 52 - 68 cm (females); weight 2: 5 — 4 - 5 kg (males) and 2.3 - 8 kg (females). The distinguishing characteristic of the Spot-nosed Monkeyis, as its name suggests, a noticeable spot of white hair on the nose. This light patch of hair contrasts strongly against a dark face. A black brow band extends around face and behind ears to form a border around the crown, which is speckled yellow and black. There are bushy white whiskers on lower cheeks, and throat has a ruff of white hair. Ears are prominent and have conspicuous fringes of white hairs. On each side of the head, a linear tuft, or “ flash, ” of white hair extends laterally below each ear. Back and flanks are agouti-brown, and underside (chest and belly) is white. Limbs are pale on inner surfaces and speckled like the body on outside lower segments. Dorsal surface oftail is also a speckled agouti-brown, becoming darker toward tip, while its undersurface is grayish-white. Scrotum is blue. The two subspecies are very similar in appearance. The “ Eastern Lesser Spot-nosed Monkey ” (C. p. petaurista) has a black brow band extending around its face and behindits ears, but the “ Western Lesser Spot-nosed Monkey ” (C. p. buettikoferi) does not. Some individual Western Lesser Spot-nosed Monkeys have bilateral patches of creamy-speckled hairs on their cheeks; however, this seems to be a variable characteristic because the cheek patch in some individuals is not obvious or is instead silvery.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFDFFF9FA006FFEFE66F358.taxon	biology_ecology	Habitat. Primary and secondary forest, riparian and gallery forest, woodland savanna, coastal bushland, plantations, and gardens. The Spot-nosed Monkey flourishes in disturbed forest, including “ farmbush ” (secondary growth and scrub on farmland), where it is often more abundant than in mature high forest. Spot-nosed Monkeys prefer thick, young secondary growth.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFDFFF9FA006FFEFE66F358.taxon	food_feeding	Food and Feeding. Major foods of the Spot-nosed Monkey include leaves, fruits, shoots, flowers, and insects. In the Tai Forest, these monkeys spend 40 % of their feeding time eating foliage (young leaves and new stems from lianas being most frequently eaten foliage items) and 34 % eating fruits (with fruit of the canopy tree Dialium aubrevillei, Fabaceae, being the most popular item offruit). Spot-nosed Monkeys also occasionally raid maize and cacao crops.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFDFFF9FA006FFEFE66F358.taxon	breeding	Breeding. There is no specific information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFDFFF9FA006FFEFE66F358.taxon	activity	Activity patterns. The Spot-nosed Monkey is highly arboreal and rarely goes to the ground. In the Tai Forest, nearly 70 % of an individual's time is spent in the forest understory. Individuals rarely use upper parts of the main forest canopy and hardly ever enter crowns of tall emergent trees. Spot-nosed Monkeys climb quadrupedally on branches, twigs, and climber stems, avoiding large boughs. In the Tai Forest, they spend 45 % of their time feeding and foraging, 26 % moving, 24 % resting, and the remainder engaging in other activities.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFDFFF9FA006FFEFE66F358.taxon	biology_ecology	Movements, Home range and Social organization. Group sizes of Spot-nosed Monkeys are 4 - 24 individuals, although most groups are relatively small, with an average of eleven individuals. Groups generally comprise a single adult male with a number of adult females and young. In Tai, there are groups with more than one adult male. Group members interact very little; females occasionally groom other females, males, and young. A group will travel 500 - 1800 m / day, with an average of c ¢. 1000 m / day. Annual home ranges of social groups are 40 - 100 ha. Home range overlap is high among groups of Spot-nosed Monkeys. Aggression by both sexes occurs when groups meet, especially at feeding trees.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFDFFF9FA006FFEFE66F358.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Least Concern on The [UCN Red Lust, including both subspecies. The Spot-nosed Monkey is listed as Class B in the African Convention on the Conservation of Nature and Natural Resources. Although it faces threats from habitat loss through deforestation and human settlement, it is evidently able to survive in variously degraded habitats. Its preference for regrowth forest over mature high forest has so far enabled it to withstand human pressures across its distribution. The hunting of Spot-nosed Monkeys is also not as severe due to its small size and its habit of hiding in thick undergrowth where it is difficult to shoot. A high visual acuity also means that individuals are often able to detect the approach of people well in advance of a threat. The Western Lesser Spot-nosed Monkey occurs in Tai Forest National Park, Monogaga Classified Forest, and Kabéoula and Gouléako II community reserves in Ivory Coast, and the Eastern Lesser Spot-nosed Monkey occurs in Digya National Park in Ghana. The Spot-nosed Monkey is a common and widespread species. Although facing threats from habitat loss and hunting, the subspecies are also well able to persist in a wide range of degraded habitats.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFCFFF9FFE669F8F708F26C.taxon	materials_examined	West Africa. Restricted by P. Grubb and colleagues in 1999 to Benin, Lama Forest.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFCFFF9FFE669F8F708F26C.taxon	discussion	C. erythrogasteris a member of the C. cephus superspecies group as defined by J. Kingdon in his 1997 Field Guide to African Mammals. It also includes C. cephus, C. petaurista, C. sclateri, C. erythrotis, and C. ascanius. C. P. Groves in his 2001 Primate Taxonomy had the same composition for his cephus species group. Two subspecies recognized.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFCFFF9FFE669F8F708F26C.taxon	distribution	Subspecies and Distribution. C. e. erythrogasterGray, 1866 — SETogo (TogodoForest) andSBenin (LamaClassifiedForestandontheNigerianborder, NofPortoNovo). C. e. pococki Grubb, Lernould & Oates, 1999 — SW & SC Nigeria.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFCFFF9FFE669F8F708F26C.taxon	description	Descriptive notes. Head-body 45 - 50 cm (males) and 40 - 45 cm (females), tail 65 - 70 cm (males) and 55 - 60 cm (females); weight 3 - 5 — 4 - 5 kg (males) and 2 - 4 kg (females). The Red-bellied Monkey is characterized by its prominent white throat — essentially a ruff of white hairs under lowerjaw that extends onto sides of neck. Another distinguishing feature is a black-edged diadem of gold-flecked black hairs on the front of the crown. The gold contrasts with solid black frontal and parietal bands. Cheeks have tufts of light yellow and black banded hairs with white whiskers. Nose patch may be white, black, or dark brown. Back and flanks are a dark agouti, red-brown color, and underside is gray or red. Arms are dark, almost black, and legs are a lighter gray. Tail has a pale, grayish-white ventral surface and a dark tip. Red-bellied Monkeys typically hold their tail in a vertical question-mark pose, with tip away from head and body. The “ Red-bellied Guenon ” (C. e. erythrogaster) has a bright rusty-red chest and underside, except groin, underside of tail base, and inner sides of thighs, which are white. Its black fronto-parietal band reaches to just behind ears. The “ Nigeria White-throated Monkey ” (C. e. pococki) has a brownish-gray chest and belly, sometimes with a slight reddish tinge. Its black fronto-parietal band reaches right around behind the crown and mayjoin to complete the circle at the back.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFCFFF9FFE669F8F708F26C.taxon	biology_ecology	Habitat. Lowland moist and semi-deciduous forest, including riverine gallery and swamp forest. Red-bellied Monkeys inhabit primary and secondary forest, although most of the forest in which they now live has been heavily modified by human activities.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFCFFF9FFE669F8F708F26C.taxon	food_feeding	Food and Feeding. Diet and feeding behavior of the Red-bellied Monkey are poorly known. Principal food items are thought to be fruits, and it also eats leaves and insects. In the Lama Forest, Benin, it favors ripe fruit, with immature fruit eaten in the dry season. It is also raids crops because of forced habitat restriction.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFCFFF9FFE669F8F708F26C.taxon	breeding	Breeding. There is no specific information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFCFFF9FFE669F8F708F26C.taxon	activity	Activity patterns. The Red-bellied Monkey prefers dense, tangled forest vegetation and spends much of its time in lower levels of the canopy and among growth in canopy gaps along rivers. Individuals are agile and use small to medium-sized supports to walk, run, and climb quadrupedally through the forest. This, coupled with their acute vision, makes them difficult to observe because they often detect people well before people are able to detect them.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFCFFF9FFE669F8F708F26C.taxon	biology_ecology	Movements, Home range and Social organization. The Red-bellied Monkey lives in social groups, although there are no accurate data on group composition. Most groups probably have 10 - 20 individuals, although groups of up to 30 individuals have been observed. They are most likely to contain a single adult male, adult females, and young. Solitary adult males have also been observed. On census walks in Lama, Benin, groups averaged 9 - 6 individuals, butit is typical not to see all group members during surveys ofthis type. No published information on movements and home range is available.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFCFFF9FFE669F8F708F26C.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Vulnerable on The IUCN Red List, including the subspecies pococki, but the nominate subspecies erythrogaster is classified as Endangered. The Red-bellied Monkeyis listed as Class A in the African Convention on the Conservation of Nature and Natural Resources. Habitat loss and hunting have led to a population decline of 30 % over the past 27 years (three generations). Their forests have been severely degraded and exploited by agricultural expansion and timber extraction. The Red-bellied Guenon has a scattered distribution, but it is relatively abundant in some areas such as Okomu National Park and in a number of forest reserves (e. g. Udo, Omo, Oluwa, and Ifon). The Red-bellied Guenon has a restricted range (less than 5000 km?) in a heavily populated area. This, coupled with intense hunting, is causing continuing population decline, and it is likely that the population of Red-bellied Guenons has declined by ¢. 50 % over the past 27 years. It persists in forests that are not suitable for cultivation because to seasonal flooding and occurs in some sacred tree groves (where hunting is taboo) and gallery forests. A protected central refuge of semi-deciduous forest in the Lama Forest of Benin (c. 2000 ha) is home to the largest remaining population of the Red-bellied Guenon. Facing similar threats, the Nigeria White-throated Monkey is believed to have declined ¢. 30 - 40 % over the past 27 years. Its populations occur in the central part of Okomu Forest Reserve in Edo State (Okomu became a Wildlife Sanctuary in 1985 and a National Park in 1999) and Omo Forest Reserve in Ogun State, which includes a 460 ha Biosphere Reserve (a larger wildlife conservation area is also planned for this locality). The Nigeria White-throated Monkey also occurs in the Niger Delta where, although the forest is degraded and not protected, hunting pressure is lower than in most other parts of southern Nigeria.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFFFFFAFF34641FFD6FF270.taxon	materials_examined	Nigeria, Benin City.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFFFFFAFF34641FFD6FF270.taxon	discussion	C. sclateri 1 s a member of the C. cephus superspecies group as defined byJ. Kingdon in his 1997 Field Guide to African Mammals. It also includes C. cephus, C. petaurista, C. erythrotis, C. ascanius, and C. erythrogaster. C. P. Groves in his 2001 Primate Taxonomy had the same composition for his cephus species group. Individuals found toward the Cross River Valley have varying tints of red on the nose and ears. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFFFFFAFF34641FFD6FF270.taxon	distribution	Distribution. SE Nigeria, in a few small, scattered populations in forest and derived savanna between the lower Niger and Cross rivers, including the Niger Delta (specifically, from W of Oguta at 5 ° 42 ° N, just E of the Niger); it occurs N as far as the Eastern Aboine River in Ebonyi State.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFFFFFAFF34641FFD6FF270.taxon	description	Descriptive notes. Head-body 45 - 55 cm (males) and 40 - 45 cm (females), tail 60 - 75 cm (males) and 45 - 60 cm (females); weight 3 — 4 - 5 kg (males) and 2.5 - 3.5 kg (females). Anterior crown of Sclater’s Monkey is brindled black and yellow, forming a diadem of gold-flecked hair, bordered posteriorly by a black band that runs from eye to ear and around back of the crown. Throat is white but does not form a distinctive ruff as seen in the Red-bellied Monkey (C. erythrogaster). There is a narrow, pale brow band, and facial skin around eyes is pale bluish or pink. Skin of muzzle, including tip of nose, is pale and pinkish (sometimes with a reddish edge). Cheeks have tufts of lemon-yellow hairs, with light yellow whiskers. Ears have a prominent fringe of white hairs forming tufts. Back, flanks, outer limb surfaces, and dorsal proximal surface are speckled gray olive-brown (“ agouti ” coloration). Ventral side and forearms are grayish. Dorsal side oftail is the same color as back, and it has a red ventral base becoming white in the middle and darkening toward tip; however, there is considerable individual variation in this feature.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFFFFFAFF34641FFD6FF270.taxon	biology_ecology	Habitat. Remnant secondary, gallery, riparian, and freshwater swamp forests. Sclater’s Monkey originally occurred in moist tropical lowland forest, but because of severe habitat degradation, it now persists in remnant habitats. It is also found in “ farmbush ” (secondary growth and scrub on farmland), and small remnant tree groves (sometimes where only scattered trees remain in villages) where it is regarded as sacred. All sites from which it is known are below 130 m above sea level.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFFFFFAFF34641FFD6FF270.taxon	food_feeding	Food and Feeding. Diets of Sclater’s Monkey are no doubt mainly fruits, along with leaves and animal prey. In villages where they are considered sacred, they eat agricultural crops (such as oil palm nuts) and steal food from houses.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFFFFFAFF34641FFD6FF270.taxon	breeding	Breeding. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFFFFFAFF34641FFD6FF270.taxon	activity	Activity patterns. Sclater’s Monkey occupies all levels of the forest canopy, but in areas where they are hunted, they spend much of their time in thick, low growth. No further information on the activity patterns is available.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFFFFFAFF34641FFD6FF270.taxon	biology_ecology	Movements, Home range and Social organization. There is limited information on the social organization of Sclater’s Monkey. Short-term observations found groups of 3 - 30 individuals. Groups probably comprise a single adult male, adult females, and young. Evidence for this comes from their vocalizations; loud calls usually come from only one location in a group (indicating the presence of a single male). The home range and ranging behavior of Sclater’s Monkey have not been studied.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFFFFFAFF34641FFD6FF270.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Vulnerable on The IUCN Red Lust. Sclater’s Monkeyis listed as Class B in the African Convention on the Conservation of Nature and Natural Resources. It has undergone a population decline exceeding 30 % over the past 27 years (three generations). Principle threats include habitat loss and degradation and hunting. Sclater’s Monkey was once feared to be extinct in the wild; until the 1980 s, no wild populations had been observed. It was known only from a single captive individual first described in 1904. In 1988, Sclater’s Monkey was found just south-west of Oguta, Im o State, and further surveys have since confirmed its presence at more than 30 sites. Most of them are in a few, small, fragmented, and degraded forests, with correspondingly small populations. Sclater’s Monkey occurs in a region with one of the densest human populations in all of Africa. Much of the original habitat has been converted to cultivated land, used for petroleum exploration, or has been destroyed by logging. The level of fragmentation is of particular concern as populations become ever more separated and isolated. Sclater’s Monkey has a slight advantage over other guenons with regard to hunting; they are small and elusive, and not a preferred or commonly caught species. Sclater’s Monkey receives some protection because of traditional beliefs in a few Igbo and Ibibio villages where it is viewed as sacred, but it does not occur in any areas that are strictly protected by governmental legislation for wildlife conservation. Sclater’s Monkeys are found in Stubbs Creek Forest Reserve in Akwa Ibom State, Upper Orashi Forest Reserve in Rivers State, and Edumanom Forest Reserve in Bayelsa State. All of these reserves contain fragmented and degraded forest, and the wildlife suffers from unregulated hunting. The continuing degradation and fragmentation of remnant habitats in southern Nigeria are serious threats to the survival of Sclater’s Monkey.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFFFFFBFA316402FC88FDA8.taxon	materials_examined	Equatorial Guinea, Bioko Island.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFFFFFBFA316402FC88FDA8.taxon	discussion	C. erythrotisis a member of the C. cephus superspecies group as defined byJ. Kingdon in his 1997 Field Guide to African Mammals. It also includes C. cephus, C. petaurista, C. sclaten, C. ascanius, and C. erythrogaster. C. P. Groves in his 2001 Primate Taxonomy had the same composition for his cephus species group. The subspecies camerunensis hybridizes with C. ¢. cephus in a small area south of the Sanaga River in Cameroon. Two subspecies recognized.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFFFFFBFA316402FC88FDA8.taxon	distribution	Subspecies and Distribution. C. e. erythrotisWaterhouse, 1838 — mountainousareasinC & SBiokoI (EquatorialGuinea). C. e. camerunensis Hayman, 1940 — SE Nigeria (E of lower Cross River) and NW Cameroon (N of lower Sanaga River), also in a small areajust S of the Sanaga River, near its mouth, between Tinaso and Lake Tisongo.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFFFFFBFA316402FC88FDA8.taxon	description	Descriptive notes. Head-body 45 - 55 cm (males) and 40 - 45 cm (females), tail 56 — 77 cm (males) and 46 - 65 cm (females); weight 3 - 5 — 4 - 5 kg (males) and 2.3 - 3.5 kg (females). The Red-eared Monkey takes its name from a fringe of rusty-red hairs on outer margin ofits ears. A more distinctive feature is its rusty-red tail. Ventrally, tail is red for most ofits length, dorsal surface is a darker brown-red, and tip is blackish. A black brow band extends to ears. Bare skin around eyes is bluish-gray, and muzzle is naked and pink. Upper cheeks have tufts of white and lemon-yellow hairs, and lower cheeks are black. There is a triangular reddish nose spot, and throatis whitish. Crown of head, back, and upper outer leg surfaces are grayish, with orange-brown speckles, becoming grayer on flanks. Lower arms are dark gray, and belly and inner surfaces of limbs are gray. Scrotum is covered in bright orange-red hairs, and the penis is pink. Vulva is light blue. In both sexes, callosities are chocolate-colored. The “ Bioko Red-eared Monkey ” (C. e. erythrotis) is smaller, with a darker pelage and shorter tail. The “ Cameroon Redeared Monkey ” (C. e. camerunensis) has a lighter colored, yellower pelage with shorter hairs. It is larger than the Bioko Red-eared Monkey and has a longertail.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFFFFFBFA316402FC88FDA8.taxon	biology_ecology	Habitat. The Bioko Red-eared Monkey occurs in primary and secondary lowland tropical and submontane moist forest near rivers. On the mainland, the Cameroon Redeared Monkey is more abundant in mature forest than young secondary forest. This may be because of competition with the Diana Monkey (C. diana), which flourishes in disturbed and edge habitats. The Cameroon Red-eared Monkey is also more abundant in wetter forests in the southern part of its distribution than in more northerly semideciduous forests. On Bioko Island (where the Diana Monkey is absent), the Bioko Red-eared Monkey is the most abundant monkey species. There, it is most commonly found in mature lowland forest, although it also occurs in secondary lowland forest and overgrown cacao plantations, and above an elevation of 2000 m on Pico Basilé in montane forest and grassland. It will also live in close proximity to people on Bioko Island.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFFFFFBFA316402FC88FDA8.taxon	food_feeding	Food and Feeding. Fruits are evidently a major component of the diets of Red-eared Monkeys, along with leaves, shoots, and arthropods. They also raid agricultural crops.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFFFFFBFA316402FC88FDA8.taxon	breeding	Breeding. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFFFFFBFA316402FC88FDA8.taxon	activity	Activity patterns. The Red-eared Monkey uses all levels of the forest canopy. On Bioko Island, they are generally seen at 5 - 25 m above the ground but sometimes in the upper canopy at 45 m. In Cameroon, they feed and travel lower in the forest canopy than other arboreal guenons. Red-eared Monkeys also go to the ground and have been seen traveling through montane grassland and on beaches on southern Bioko Island.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFFFFFBFA316402FC88FDA8.taxon	biology_ecology	Movements, Home range and Social organization. Groups of 8 - 12 Red-eared Monkeys are typical on south-eastern Bioko, whereas they have 4 - 30 individuals in south-western Cameroon. Groups have a single male, adult females, and young. No published information is available on their home range or ranging behavior.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFFFFFBFA316402FC88FDA8.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Vulnerable on The [UCN Red List, including both subspecies. The Red-eared Monkeyis listed as Class B in the African Convention on the Conservation of Nature and Natural Resources. It is believed to have declined in numbers by 30 % over the past 27 years (three generations) because of hunting and habitat loss. The bushmeattrade is a significant threat to Redeared Monkeys, and it is the most common monkey in bushmeat markets in Malabo on Bioko Island. A survey counted 622 Red-eared Monkey carcasses for sale in the Malabo market in March — October 1991. An estimated 1025 Red-eared Monkey carcasses are sold annually at three markets in and around Cameroon’s Korup National Park. Despite this, Red-eared Monkeys still manage to persist in relatively high abundance in some areas. They are elusive and can be hard to detect, and they are reasonably adept at evading capture by bushmeat hunters. This has enabled them to survive over a large area in much of their historic distribution. Although also threatened by deforestation through timber extraction and conversion of forest to agricultural land, they are the most common primate on Bioko Island. In 2006, the Bioko Red-eared Monkey was estimated to number ¢. 21,000 individuals, down from more than 30,000 individuals in 1986. The Cameroon Red-eared Monkey is believed to have undergone a decline of c. 30 % over the past 27 years. Red-eared Monkeys are protected by national legislation in Cameroon and Nigeria, and they can be found in several protected areas, including Korup National Park and several forest reserves in Cameroon, Cross River National Park in Nigeria, and Basilé National Park and Southern Island Scientific Reserve on Bioko Island. Despite such protection, hunting is often as severe inside protected areas as it 1 s outside of them.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFEFFFBFFEE678CFAA5F273.taxon	materials_examined	America (= West Africa).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFEFFFBFFEE678CFAA5F273.taxon	discussion	C. cephus is a member of the C. cephus superspecies group as defined byJ. Kingdon in his 1997 Field Guide to African Mammals. It also includes C. petaurista, C. sclateri, C. erythrotis, C. ascanius, and C. erythrogaster. C. P. Groves in his 2001 Primate Taxonomy had the same composition for his cephus species group. Albinism has been reported. The subspecies cephus hybridizes with C. e. camerunensis in a small area just to the south of the Sanaga River nearits mouth, between Tinaso and Lake Tisongo. Three subspecies recognized.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFEFFFBFFEE678CFAA5F273.taxon	distribution	Subspecies and Distribution. C. c. cephusLinnaeus, 1758 — SCameroon (SofSanagaRiver), extremeSWofCentralAfricanRepublic (EtotheMambéréRiver), EquatorialGuinea, NEGabon, mostoftheRepublicoftheCongo, andNWAngola. C. c. cephodesPocock, 1907 — WGabonandWRepublicoftheCongo, incoastalregionsbetweentheOgoouéandKouilou-Niaririvers. C. c. ngottoensis Colyn, 1999 — SW Central African Republic (S of the Mambéré-Kadéi Prefecture and Bangui), N Republic of the Congo (E to the Ubangi River), possibly in NW DR Congo.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFEFFFBFFEE678CFAA5F273.taxon	description	Descriptive notes. Head-body 50 - 58 cm (males) and 44 - 50 cm (females), tail 65 — 99 cm (males) and 60 - 76 cm (females); weight 3.8 - 5 kg (males) and 2 — 4 kg (females). The Mustached Monkey has a distinctive, chevron-shaped, white moustache that contrasts strongly with yellowish-orange cheek tufts and bare, dark blue skin of face. Coat is speckled reddish-brown, becoming uniformly dark gray toward extremities. Throat and belly are whitish gray. The “ Red-tailed Mustached Monkey ” (C. c. cephus) is a small guenon, and its tail is much longer than its body. In the Red-tailed Mustached Monkey and the “ White-nosed Mustached Monkey ” (C. ¢. ngottoensis), tails are mostly blackishgray, except for the lower part, which is red. Tail of the “ Gray-tailed Mustached Monkey ” (C. c. cephodes) is brown and gray.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFEFFFBFFEE678CFAA5F273.taxon	biology_ecology	Habitat. Lowland tropical, gallery, and riparian forest. The Mustached Monkey will also tolerate secondary habitat types where scrub or forest is in the process of growing back after logging, fire, or some other major disturbance.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFEFFFBFFEE678CFAA5F273.taxon	food_feeding	Food and Feeding. The Mustached Monkey is primarily frugivorous. Fleshy ripe fruit makes up ¢. 80 % ofits diet. It also seems to be relatively dependent upon nuts of oil palms because individuals are rare or totally absent in areas lacking such trees. Seeds, leaves, and arthropods are important constituents of the diet, and they occasionally eat eggs and fledglings.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFEFFFBFFEE678CFAA5F273.taxon	breeding	Breeding. Although breeding may occur throughout the year, births in Gabon occur in December — February, peaking in January. Single young are born after a gestation of 5 - 6 months.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFEFFFBFFEE678CFAA5F273.taxon	activity	Activity patterns. Mustached Monkeys prefer the lower strata of the forest. They spend c. 60 % of their time below a height of 15 m above the ground. They are one of the smaller guenons and generally forage where there is dense forest cover.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFEFFFBFFEE678CFAA5F273.taxon	biology_ecology	Movements, Home range and Social organization. Groups of Mustached Monkeys contain 4 - 35 individuals, commonly a single male, multiple females, and their young. No published information on movements or home rangesis available.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFEFFFBFFEE678CFAA5F273.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Least Concern on The [IUCN Red List, including the subspecies cephus. The subspecies ngottoensis is classified as Data Deficient, and cephodes has not been assessed. The Mustached Monkeyis listed as Class B in the African Convention on the Conservation of Nature and Natural Resources. The Mustached Monkey is relatively common and widespread and is not currently facing any major threats to warrantlisting in a category of threat. Its main threats include habitat loss through deforestation and hunting for meat. As deforestation rates and the bushmeat trade continue to grow, it is possible that it will undergo substantial decline. The Mustached Monkey is able to adapt to secondary habitat and is also present in a number of protected areas in Gabon and southern Cameroon. The Gray-tailed Mustached Monkey is found in a number of protected areas and is therefore safeguarded to some extent from habitat loss. The White-nosed Mustached Monkey has not been discovered in any existing protected areas within its range. It is, however, found within the boundaries of the proposed Mbaere-Bodingue Park in the Central African Republic, which will hopefully provide a refuge in the future.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFEFF84FAE86402F8BCF9D0.taxon	materials_examined	no type locality specified. Restricted by A. B. Machado in 1969 to north-west Angola, by the lower Congo River.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFEFF84FAE86402F8BCF9D0.taxon	discussion	C. ascanius was previously associated with C. petaurista, C. erythrogaster, and C. nictitans, all spot-nosed monkeys. It is now agreed that the cephus species group should include all lesser spot-nosed monkeys. Genetic studies suggest a relatively recent divergence from other cephus monkeys. This polytypic species requires further investigation to assess the taxonomic distinctiveness of the geographic types. P. Grubb and colleagues, in their 2003 review, raised doubts about the subspecific status of atrinasus, suggesting that more research was necessary to confirm its taxonomic validity. C. P. Groves in his 2001 Primate Taxonomy recognized five subspecies but considered the possibility of raising schmidti to a full species because it lacks the temporal whorl found in the other subspecies. C. ascanus is reported to hybridize with C. mitis stuhlmanni, C. doggetti, and C. denti. There is a transitional zone of C. a. ascanius x C. a. atrinasus where their distributions meet in north-central Angola. Intermediates of C. a. whitesidei and C. a. katangae occur between the Lualaba and Lomami rivers. C. a. atrinasus x C. a. katangae hybrids are reported from northern Angola. Five subspecies recognized.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFEFF84FAE86402F8BCF9D0.taxon	distribution	Subspecies and Distribution. C. a. ascaniusAudebert, 1799 — SWDRCongo (SoftheCongoandKasairivers) andNAngola (Sboundaries ¢. 8 ° SandElimitatc. 15 ° E); furtherEisanapparenthybridzonewiththesubspeciesatrinasusbuttheboundariesareuncertain. C. a. atrinasusMachado, 1965 — NEAngola, knownonlyfromasmallareaaroundthetypelocalityintheNWcorneroftheLundaDistrict, aroundtheWambaRiver; ithasbeenmappedfromc. 8 ° - 9 ° §, c. 18 ° - 18 ° 30 ’ E. C. a. katangaeL. onnberg, 1919 — SDRCongo (mainlybetweentheKasaiandLualabarivers), NWZambia, andNAngola (to 11 ° S, 17 ° E, exceptforthesmallsubspeciesatrinasusenclave). C. a. schmidtiMatschie, 1892 — DRCongo (EoftheLualabaRiver) extendingintoUganda, Rwanda, WBurundi, SWKenya, andWTanzania; NoftheCongoRiver, itextendsNandWoftheUbangiRiverintotheCentralAfricanRepublicandSSouthSudan. C. a. whiteside: Thomas, 1909 — DR Congo, between the Congo and the Kasai-Sankuru rivers. Its border with the subspecies katangaeis a little south of the Sankuru, with intermediates occurring between the Lualaba and Lomami rivers.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFEFF84FAE86402F8BCF9D0.taxon	description	Descriptive notes. Head-body 43 - 51 - 56 cm (males) and 34 - 48.5 cm (females), tail 60 - 92 cm (males) and 54 - 79 cm (females); weight 3 - 6 kg (males) and 1.8 - 4 kg (females). Dorsal coat of the Red-tailed Monkey is agouti-brown, with light (white, cream, or gray) on underside and inner side of limbs. Hair bases on body are gray, with two to four alternating red / black bands, the first pair separated from the light base by an indistinct black band; subterminal black band is longer than the rest. At least part of tail is red, with a black tip. Hands and feet are dark. There is a conspicuous heart-shaped nose spot that is white, yellow, red, or black, depending upon the subspecies. In most subspecies, there is a whorl in front of ears, from which white cheek whiskers radiate in a spiral-like pattern, almost vertical in front, and sweeping down and back below. Whiskers are bordered inferiorly by a black band, curving up medially alongside the nose spot. Circumocular bare skin is slate blue or black. There is also a blackjaw band. In the “ Black-cheeked Red-tailed Monkey ” (C. a. ascanius), nose spot and temporal whorls are white. Ear tufts are bright orange-red, and cheeks are black, with black frontal and temporal bands being notably wide. Periocular skin is light blue. Its underside is gray, extending under base oftail. Tail tip is darker. Hairs are gray at their bases, with four pairs of alternating red and black bands. The “ Black-nosed Red-tailed Monkey ” (C. a. atrinasus) is similar to the Black-cheeked Red-tailed Monkey, but it has a black nose spot, blue circumocular rings, and black maxilla. White hairs emerging from its cheek whorl are short. Its cheek whiskers are white above, black in the middle due to lateral extension of the black maxillary zone, and gray below, extending to its throat. The “ Yellow-nosed Red-tailed Monkey ” (C. a. whitesidei) has a creamy yellow to deep orange nose spot and reddish ear tufts. Its black frontal band is narrow, with a reduced inferior lateral black band. Its whitish underside extends to underside of tail. Dorsal surface of tail is dark, with the distal one-third red. Its cheek whiskers do not have as much of a lateral flare as those of Black-cheeked and Black-nosed red-tailed monkeys. Whiskers are white superiorly, agouti-brown inferiorly, and gray at the throat. Periocular skin is almost black. In the “ Katanga Red-tailed Monkey ” (C. a. katangae), nose spot and temporal whorls are white, and skin around eyes is violet purple. Its black frontal band is narrow, and its black temporal band is reduced. Cheeks have a reduced black area, and ear tufts are reddish. Tail is very dark above with a blackish, rather than red, distal portion. Hairs differ from other subspecies in that they are white and not gray at the base with the same alternating bands of red and black. “ Schmidt's Red-tailed Monkey ” (C. a. schmidti) has a distinctly different face from other subspecies. Skin around eyes is a dark blue to nearly black. Nose spot and ear tufts are white. Frontal band and inferior lateral black bands are narrow. Cheeks have an extensive whitish area that extends back to partially cover ears, margined with black below. A temporal whorl is distinctly lacking in this subspecies. Its appearance is so distinctive that is has prompted debate regarding the possibility of considering it a full species. There are even reports of individuals in the southern part ofits distribution in DR Congo that have dark gray legs, sometimes extending to the back.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFEFF84FAE86402F8BCF9D0.taxon	biology_ecology	Habitat. A wide range of lowland, submontane, and montane forests, both dry and moist, including tropical forest, swamp forest, riparian and gallery forest, deciduous forest, forest mosaics, and Acacia (Fabaceae) woodland. Red-tailed monkeys occur in primary and disturbed secondary growth forests. The more lowland, gallery rainforest habitats are preferred by Black-cheeked Red-tailed Monkeys. Higher elevations and dry forests are inhabited by Schmidt's Red-tailed Monkey. They are reported in Brachystegia (Fabaceae) and undifferentiated woodland and on cultivated subsistence farmland and tree plantations. They are usually found near rivers and range from zero to 2500 m above sea level. Black-cheeked Red-tailed Monkeys occur at average elevations of 800 m, and Schmidt’s Red-tailed Monkey at 400 - 2500 m.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFEFF84FAE86402F8BCF9D0.taxon	food_feeding	Food and Feeding. Diets of Red-tailed Monkeys contain at least 60 % fruit, along with seeds, young and mature leaves, flowers, nectar, and sometimes bark and gum. Insects can make up 30 % of the diet. Other animal matter consumed includes birds and small reptiles. They hunt African green-pigeons (7 reron calva) in the Kalinzu Forest in Uganda. They have been reported to raid banana, mango, guava, and maize crops near Kibale Forest National Park, Uganda.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFEFF84FAE86402F8BCF9D0.taxon	breeding	Breeding. Female Red-tailed Monkeys have a menstrual cycle of ¢. 30 days. The gestation period is 5 - 6 months, and singletons are the norm. Mating generally occurs throughout the year, with some seasonality apparent in some regions correlated with rainfall and food availability. Exclusive maternal care and allomothering are practiced. Infanticide has been reported. Females reach sexual maturity at c. 4 years of age and males at six years. Males emigrate from natal groups. The life span is c. 28 years.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFEFF84FAE86402F8BCF9D0.taxon	activity	Activity patterns. Red-tailed Monkeys are diurnal and arboreal. They spend 60 - 90 % of their time at heights of 5 - 20 m above the ground, using an average forest height of c. 12 m. Locomotion is quadrupedal, with climbing and leaping.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFEFF84FAE86402F8BCF9D0.taxon	biology_ecology	Movements, Home range and Social organization. Red-tailed Monkeys live in hierarchical, male-dominant, unimale-multifemale social groups. Average group size is 30 - 35 individuals. A study carried out at Ngogo in Kibale Forest National Park, Uganda, reported the maximum supportable group size of Schmidt's Red-tailed Monkey to be c. 50 individuals, roughly 50 % larger than the average group size of the species. This study reported two occasions, and a possible third, of group fission when a group reached 50 individuals. Studies of Schmidt's Red-tailed Monkey reported daily movements of 1400 - 1900 m. Home ranges are 15 - 68 ha.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFFEFF84FAE86402F8BCF9D0.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Least Concern on The [UCN Red List, including the subspecies schmidti. The subspecies atrinasus is classified as Data Deficient, and katangae, whitesidei, and the nominate subspecies ascanius have not been assessed. The Red-tailed Monkeyis listed as Class B in the African Convention on the Conservation of Nature and Natural Resources. Because of its occurrence in a wide variety of forest habitats, it has no major threats. Two subspecies have been evaluated. Schmidt's Red-tailed Monkey has a wide distribution in a variety of habitats, and although it is locally threatened by hunting and habitat loss and degradation, there are apparently are no threats great enough to result in a range-wide decline that would warrant listing in a threatened category. The Black-nosed Red-tailed Monkey is known only from data collected on nine individuals from the north-west Lunda District of north-eastern Angola. The remaining three subspecies have not been assessed. In general, threats to Red-tailed Monkeys include habitat encroachment from agriculture and forest exploitation. They are locally threatened by habitat loss and hunting, but they are probably able to tolerate a degree of hunting. They are hunted for food and fur (its tail fur was formerly used to cover bows). It is often killed as a crop pest, and it is used in biomedical research. Red-tailed Monkeys occur in at least 15 protected areas, including Kahuzi-Biéga and Salonga national parks, Tayna Gorilla Reserve, and Okapi Wildlife Reserve in DR Congo; Nyungwe Forest and Volcanoes national parks in Rwanda; Gombe Stream and Mahale Mountains national parks in Tanzania; Bwindi Impenetrable, Kibale Forest, and Ruwenzori national parks and Budongo, Itwara and Mabira forest reserves in Uganda; and Kakamega Nature Reserve in Kenya.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF83FF86FF356405F938F72E.taxon	materials_examined	Guinea. Restricted by C. P. Groves in 2005 to Equatorial Guinea, Benito River.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF83FF86FF356405F938F72E.taxon	discussion	In his 1997 Field Guide to African Mammals, J. Kingdon aligned C. nictitans with C. mitis, comprising the “ gentle monkey, ” C. nictitans superspecies group. C. matis in this arrangement included four clusters: Mitis monkeys, white-throated monkeys (including albogularis), silver monkeys (including doggetti and kandti), and a cluster that includes the forms stuhlmanna, elgonis, and boutourlinii. Each cluster circumscribed a potential species. This arrangement was followed in The Mammals ofAfrica (Volume 2), edited by T. Butynski and colleagues (published in 2013). The third contemporary arrangement for this group was published by P. Grubb and colleagues in 2003. It is similar to that of Kingdon with just two species in the nictitans species group: C. nictitans and C. mitis (including subspecies albogularis, doggetti, and kandti). C. P. Groves in his 2001 Primate Taxonomy placed C. nictitans as one of five species that made up his C. mitis species group; the others were C. matis, C. doggetti, C. kandti, and C. albogularis. Groves recognized just two subspecies of C. nictitans: C. n. nictitans and C. n. martini. Along with those two subspecies, three additional subspecies are recognized here, following the taxonomy in J. Oates’s Primates of West Africa, published in 2011: insolitus, ludio, and stampflii (considered by Groves to be synonyms of C. n. martini). Five subspecies recognized.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF83FF86FF356405F938F72E.taxon	distribution	Subspecies and Distribution. C. n. nictitansLinnaeus, 1766 — Cameroon (extendingNoftheSanagaRiverintopartsoftheCameroonHighlands), CentralAfricanRepublic, EquatorialGuinea, NWDRCongo, Gabon, andRepublicoftheCongo; possiblyinNWAngola. C. n. insolitusElliot, 1909 — C & WNigeria. C. n. ludioGray, 1849 — SENigeriaandNWCameroon (includingMountCameroon). C. n. martiniWaterhouse, 1838 — BiokoI (EquatorialGuinea). C. n. stampfliiJentink, 1888 — N Liberia and W Ivory Coast.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF83FF86FF356405F938F72E.taxon	description	Descriptive notes. Head-body 55 - 70 cm (males) and 43 - 53 cm (females), tail 68 100 cm (males) and 56 - 76 cm (females); weight 5.5 - 8 kg (males) and 2.7 - 5 kg (females). The Putty-nosed Monkey is a large, long-tailed guenon. Its most characteristic feature, as its name suggests, is a bulbous white nose covered by an oval-shaped mat of very short, whitish hairs. This feature stands out in stark contrast to the otherwise black face and overall dark color of the body. Crown, shoulders, back, and outer legs are deep gray, speckled with olive-yellow. Outer arms, distal part of tail, hands, and feet are black. Eyes are a striking reddish-brown. The “ Bioko Putty-nosed Monkey ” (C. m. martind) is typically smaller than the other subspecies, with longer hair and a generally darker appearance. Its upper back has fewer lighter speckles than other subspecies, and its upper chest is dirty white or grayish-white. The “ Eastern Putty-nosed Monkey ” (C. n. nictitans) has a more olive-gray appearance. Fur on its head and back is black, tinged with yellow, and its underside is a dark gray. In some individuals, there are a few red hairs in the perineal region. Its lower legs are gray. “ Stampfli’s Puttynosed Monkey ” (C. n. stampflii) has creamy white hair on its chest and abdomen, and its back and tail are lighter than in the other subspecies. Its tail is speckled gray, with a black tip. The “ Nigerian Putty-nosed Monkey ” (C. n. insolitus) has a creamy-white chest and throat extending along inner sides of upper arms to its elbows. Its lower legs are dark gray to black, and its abdomen is gray. The “ Red-rumped Putty-nosed Monkey ” (C. n. ludio) has a pure white chest, with white hair extending along its inner arms to its elbows (as in the Nigerian Putty-nosed Monkey). It has reddish brown hairs on its inner thighs and below base oftail (perineum). Its lower legs are gray.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF83FF86FF356405F938F72E.taxon	biology_ecology	Habitat. Primary and older secondary lowland and montane forest, rainforest, seasonally flooded forest, and riparian forest. The Putty-nosed Monkey is less common in swamp forest. In Liberia and Ivory Coast, Stampfli’s Putty-nosed Monkey inhabits dry semi-deciduous forest to the north of the coastal moist forest zone, although this is largely thought to be the result of competitive exclusion by the Diana Monkey (C. diana) in moister habitats. In Nigeria and Cameroon, where the Putty-nosed Monkey has a widespread distribution, it shows preferences for moist and semi-deciduous forests. Here, individuals inhabit gallery forests in the forest-savanna mosaic zone north of the forest zone, and submontane forests.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF83FF86FF356405F938F72E.taxon	food_feeding	Food and Feeding. The Putty-nosed Monkey eats fruits and seeds, supplemented with leaves, gums, flowers, and arthropods (mainly ants and caterpillars). In Tai Forest, fruits made up 59 % of the diet and invertebrates 31 %. Fruits of Dialium aubrevillei (Fabaceae) and Sacoglottis gabonensis (Humiriaceae) were the most popular. Stomach contents of Eastern Putty-nosed Monkeys (killed by hunters in north-eastern Gabon) contained an average of 81 % fruits and seeds (by dry weight), 28 % leaves, and 8 % animal matter (especially caterpillars). The Putty-nosed Monkey is able to adapt well to the dry period in Gabon (May-August) by relying on leaves. This may explain how it is able to exploit relatively dry forest despite preferring moist forest where fruits and insects are more abundant. The Putty-nosed Monkey occasionally raids crops, and individuals have been reported to steal chickens.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF83FF86FF356405F938F72E.taxon	breeding	Breeding. Birth season of the Putty-nosed Monkey is December — April. A single young is born after a 5 - 6 month gestation. The annual birth rate is 0 - 41 young / year. Males reach sexual maturity at 5 - 7 years, females at c. 4 years.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF83FF86FF356405F938F72E.taxon	activity	Activity patterns. The Putty-nosed Monkey mostly occupies the middle and upper forest canopies. In western Cameroon, it uses higher levels of the forest than all other sympatric guenons. In north-eastern Gabon, most observations of Eastern Putty-nosed Monkeys were at heights of more than 20 m. In south-eastern Bioko, the Bioko Puttynosed Monkey forages and travels at heights of 10 - 30 m and is the monkey species seen most frequently at heights of 20 m or more. In the Tai Forest, Stampfli’s Puttynosed Monkey spends 75 % of its time in the main canopy and in emergents (i. e. above 15 m). It probably uses upper canopy levels more often than other guenons because of direct competitive exclusion from the Diana Monkey, also a canopy-dweller. The Putty-nosed Monkey's locomotory repertoire includes walking quadrupedally, climbing, leaping, and running. Because of their use of the uppermost levels of the canopy, individuals move around using large branches much more often than other guenons.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF83FF86FF356405F938F72E.taxon	biology_ecology	Movements, Home range and Social organization. Putty-nosed Monkeys generally live in groups of 12 - 30 individuals, with a single adult male, adult females, and young. A high level of affiliative interaction between group members is common, mostly involving grooming among females. Within-group agonistic interactions are infrequent. In Tai Forest, a group of Stampfli’s Putty-nosed Monkey used a home range of 93 ha over a 7 month period. In north-eastern Gabon, groups of Eastern Putty-nosed Monkeys have home ranges of 55 - 80 ha and daily movements of ¢. 1500 m.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF83FF86FF356405F938F72E.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Least Concern on The [UCN Red List, including the nominate subspecies nictitans, but the subspecies martini is classified as Vulnerable. Subspecies insolitus, ludio, and stampflii have not been assessed. The Putty-nosed Monkey is listed as Class B in the African Convention on the Conservation of Nature and Natural Resources. The Bioko Putty-nosed Monkey underwent a 50 % decline in its population between 1986 and 2006 because of habitat loss and hunting pressure. The Putty-nosed Monkey is threatened, especially in the western part of its distribution (Stampfli’s Putty-nosed Monkey), by habitat loss through deforestation for timber and agricultural land. There, it is also heavily hunted for meat, and it is occasionally shot as an agricultural pest. In Liberia and western Ivory Coast, competition with the Diana Monkey further combines with habitat conversion and hunting to make Stampfli’s Putty-nosed Monkeys scarce. In Ivory Coast, it occurs in Tai Forest National Park. The Putty-nosed Monkey remains reasonably common, however, in the central and eastern parts of its distribution. At Korup National Park in Cameroon, for example, its population seems to hold up in the face of heavy hunting pressure better than those of other forest monkeys. Nevertheless, the Putty-nosed Monkey is less resistant to forest disturbance and consequently is now rare in many Nigerian forest reserves that have been subjected to heavy logging, farming, and hunting. It occurs in a number of protected areas including Monte Alen National Park in Equatorial Guinea and Mbam-Djerem National Park in Cameroon.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF83FF87FA306C07F5B3F7A7.taxon	materials_examined	Guinea, but possibly Angola according to P. Napier in 1981.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF83FF87FA306C07F5B3F7A7.taxon	discussion	The taxonomy of the highly polytypic “ mitis ” group of guenonsis still unsettled and varies according to author. In his 1997 Field Guide to African Mammals J. Kingdon aligned C. nictitans with C. mutis comprising a “ gentle monkey, ” C. nictitans superspecies group. C. mutis in this arrangement included four clusters: Mitis monkeys, white-throated monkeys (includes albogularis), silver monkeys (includes doggetti and kandti), and a cluster that includes the forms stuhlmanni, elgonis, and boutourlini — 22 taxa in all. This arrangement was followed in The Mammals ofAfrica (Volume 2) edited by T. Butynski and colleagues (published in 2013). P. Grubb and colleagues in 2003 also used the arrangement of Kingdon with just two species in the C. nictitans group: C. nictitans (with two subspecies) and C. mutis (with 16 subspecies, including albogularis, doggetti, and kandti). C. P. Groves in his 2001 Primate Taxonomy placed C. mutis (with seven subspecies) as one of five species that make up his C. mitis species group, the other species being C. nictitans (with five subspecies), C. doggetti, C. kandti, and C. albogularis (with twelve subspecies). Natural and anthropogenic hybridizations have been reported: C. m. stuhlmanni x C. m. kolbi (natural) and C. m. stuhlmanni x C. ascanius schmadti, C. m. stuhlmanni x C. albogularis, C. m. stuhlmannii x C. m. schoutedeni, C. m. stuhlmannii x C. m. opusthostictus, C. m. stuhlmanni x C. m. kandti, C. m. stuhlmannii x C. doggetti, C. m. heymansi x C. m. opisthostictus (anthropogenic). Seven subspecies recognized.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF83FF87FA306C07F5B3F7A7.taxon	distribution	Subspecies and Distribution. C. m. mitisWolf, 1822 — WAngola. C. m. boutourliniiGiglioli, 1887 — SWEthiopia, fromLakeTanaSalongtheWsideoftheEthiopianRifttoNofLakeTurkana. C. m. elgonisLonnberg, 1919 — WKenya (MtElgon). IntermediatesbetweenthesubspecieselgonisandstuhlmanniiarefoundinKakamegaForestandontheeasternflanksoftheRuwenzoriMts. C. m. heymansiColyn & Verheyen, 1987 — EDRCongo (betweentheLomaniandLualabarivers, reachingtoc. 2 ° S, andextendingslightlytotheWoftheLomaniRiver). C. m. opusthostictusSclater, 1894 — SEDRCongo (fromc. 6 ° NontheWbankoftheLualabaRiverStoKatangaProvince, andWtotheLakeTanganyika), NZambia (WoftheLuangwaRiver), andEAngola. C. m. schoutedeniSchwarz, 1928 — EDRCongo, typicallyonIdjwiandShushuislandsinLakeKivu; onthemainland, specimensarefromtheWVirungaVolcanoesandasfarSWasBobandana, althoughthesestronglytendtowardthesubspeciesstuhlmanni. C. m. stuhlmanni Matschie, 1893 — S South Sudan (Didinga Hills and Imatong Mts), Uganda (N in Bunyoro, E in Tororo, and in the W), W Kenya (W of the Rift Valley, including Mt Elgon and Kakamega Forest), and NE DR Congo, from the region between the Uele and the Congo rivers, from the Itimbiri River E to the Ituri and Semliki Forests, and S to the LLualaba River c. 6 ° S.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF83FF87FA306C07F5B3F7A7.taxon	description	Descriptive notes. Head — body 46 - 71 cm (males) and 39 - 59 cm (females), tail 60 — 95 cm (males) and 49 - 88 cm (females); weight 5.9 - 9 kg (males) and 2.7 - 5.5 kg (females). Males are much larger than females. The Blue Monkey is a large, variable species, characterized by its “ diadem ” of dense pale (usually white) fur on the forehead. Upper body is speckled blue-gray, with dark blackish limbs (particularly arms) and a broad band joining them across the scapular region. Nape and crown are dark gray or black, with a pale diadem in front and long, rounded, pale cheek whiskers and short chin hairs. Tail is carried high in a slight downward curve. The “ Pluto Monkey ” (C. m. matis) 1 s dark gray above, with white-gray speckling; black limbs, underside, and proximal half oftail; dark crown and nape, banded in agouti and often continuing as a broad, black interscapular band in sharp contrast to frontal band, centrally broad and narrowing laterally; and dark gray cheek whiskers, with alternating black and white bands. Its hairs are gray at the base, with four alternating pairs of white and black bands. The “ Lomami River Blue Monkey ” (C. m. heymanst) has a blue-gray nape, crown, and shoulders, with a lighter underside than upper side; a contrasting narrow white frontal diadem; a crown that is annulated and continuous with the similarly colored mantle; and black arms. In “ Schouteden’s Blue Monkey ” (C. m. schoutedeni), pelage is long and thick, distinguished from “ Stuhlmann’s Blue Monkey ” (C. m. stuhlmanni) by its pale gray-olive color; crown, nape, and terminal half of tail are black; and it has a white-speckled frontal band. Stuhlmann’s Blue Monkey has a black crown with contrasting diadem of even width across its forehead; it is usually gray flecked with black; legs, sometimes partially black, are darker than mantle; and underside is lighter than upper side. Hairs on its body have three to five white or yellowish-white / black band pairs. There are differences according to elevation, with a blackish interorbital region in mountains west of Lake Victoria, rare in those from lower elevations. High-elevation specimens also have more developed cheek whiskers and a generally white zone from chin to nose. In the mountains of the Semliki Valley, ear tufts, which are ocher in more westerly populations, are usually white. The “ Elgon Blue Monkey ” (C. m. elgonis) is distinguished from Stuhlmann’s Blue Monkey by lack of ear tufts, very dark legs, black ventral color, and black collar on the neck. “ Boutourlini’s Blue Monkey ” (C. m. boutourlinu) differs from Stuhlmann’s Blue Monkey in that light bands of hair are yellow rather than white, with either four or five band pairs; frontal band blends with crown; and arms, legs, underside, and terminal three-quarters of tail are black. Pelage has been described as intermediate between that of Stuhlmann’s Blue Monkey and the “ Rump-spotted Blue Monkey ” (C. m. opisthostictus). The Rump-spotted Blue Monkeyis a light gray on body and legs, darker on crown and nape; arms, underside, and terminal half to three-quarters oftail are black; tail is sometimes reddened below, especially in juveniles; and there are three white / black band pairs.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF83FF87FA306C07F5B3F7A7.taxon	biology_ecology	Habitat. A variety of forests including gallery forests in wooded savanna (especially the Pluto Monkey and Stuhlmann’s Blue Monkey), lowland evergreen rainforest, riparian, montane, semi-deciduous, coniferous, coastal, swamp, secondary growth forest, and dry scrubland. Elevational range is 750 - 3300 m. The Elgon Blue Monkey is found up to 3800 m in the Rwenzori Mountains.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF83FF87FA306C07F5B3F7A7.taxon	food_feeding	Food and Feeding. Blue Monkeys have been described as dietary generalists. In lowland-type rainforest near the Equator, they are described as predominantly frugivorous, eating both ripe and unripe fruit. In regions that experience pronounced seasonality, they are reported to have more varied diets that include young and mature leaves, buds, flowers, shoots, nectar, gum, seeds, lichen, insects, and other invertebrates (primarily arthropods). A study of four groups of Blue Monkeys at Kanyawara indicated a balance of leaves (c. 33 %), fruit (c. 28 %), and insects (c. 38 %), with flowers contributing ¢. 3 - 8 %.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF83FF87FA306C07F5B3F7A7.taxon	breeding	Breeding. Sexual maturity is reached at 4 - 6 years of age. The ovarian cycle is ¢. 30 days. The gestation period is 5 - 6 months, after which usually a single infant is born. Infants are cared for by the mother and other females in a group (allomothering). Infanticide has been reported. Individuals live for 20 years, on average, but are known to live longer.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF83FF87FA306C07F5B3F7A7.taxon	activity	Activity patterns. Blue Monkeys are diurnal and arboreal. Locomotion is mainly quadrupedal. Between 13 % and 35 % of their time is spent climbing, with vertical clinging and leaping reported at ¢. 3 %. More than 50 % percent of their time is spent at heights of 10 - 20 m in the forest canopy. About 40 % oftheir time is spent at heights of 5 - 10 m, with 3 % spent between the ground and 5 m and 3 % on the ground.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF83FF87FA306C07F5B3F7A7.taxon	biology_ecology	Movements, Home range and Social organization. Home ranges vary in size according to habitat type and are c. 20 - 300 ha. Daily movement averages 1300 m. Average group size is 18 individuals. Blue Monkeys live in patriarchal, unimale-multifemale groups. Mating is polygynous. Males disperse from their natal groups, and females are philopatric.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF83FF87FA306C07F5B3F7A7.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Least Concern on The IUCN Red List, including the subspecies opisthostictus and stuhlmanni. Subspecies schoutedeni is listed as Critically Endangered, boutourlinii as Vulnerable, heymansi as Near Threatened, and mutis as Data Deficient. The subspecies elgonis has not been assessed. The Blue Monkey is listed as Class B in the African Convention on the Conservation of Nature and Natural Resources. It is generally common throughout its distribution, but it is threatened to some degree by deforestation, habitat fragmentation, and hunting for food and traditional medicine in some places. Some subspecies with restricted distributions are threatened to varying degrees. There is no recent information on the status of the Pluto Monkey. In his 1997 field guide, J. Kingdon wrote that it was possibly extinct due to its having a small distribution in an area densely populated with humans. It is a priority for further survey work. The Lomami River Blue Monkey is threatened by commercial bushmeat hunting and an expanding bushmeat market. It is limited to forests between the Lomami and Lualaba rivers — highly accessible from Kisangani. Its population status is not known, butit is expected to have experienced declines in the order of 20 - 25 % over the past 27 years (three generations) as a result of the bushmeat trade. Recent surveys of Schouteden’s Blue Monkey on the two islands where it is native have indicated that very little natural forest remains and deforestation was ongoing. No primate sightings were reported during those surveys. If Schouteden’s Blue Monkey does survive on these islands, it is likely that its numbers are very low; the status of a mainland subpopulation is questionable because it is unclear that one ever existed there. The distribution of Stuhlmann’s Blue Monkey is largely continuous in central Africa, although it is disjunct in its eastern parts. At present, the population is not believed to be declining at a rate that would warrant threatened status, although it is suffering from hunting, deforestation, and habitat degradation in some areas. Boutourlini’s Blue Monkey is restricted to tropical deciduous and riparian forests, and these habitats are threatened by extensive and uncontrolled logging for timber and agriculture. Given the extent of habitat loss, it may qualify for a higher threatened status. The Rump-spotted Blue Monkey has an extensive distribution and is widely distributed and locally common. Although it is decreasing because of habitat loss and degradation and hunting, these impacts are not believed to be resulting in a decline that would warrant listing in a threatened category. The Blue Monkey occurs in numerous protected areas acrossits distribution.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF82FF80FAEA6DB0FDFDF9CC.taxon	materials_examined	south-western Ankole, Uganda.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF82FF80FAEA6DB0FDFDF9CC.taxon	discussion	C. doggetti 1 s a member of the mitis species group of guenons, but its taxonomy remains unsettled and varies according to author. J. Kingdon in his 1997 Field Guide to African Mammals, P. Grubb and colleagues in their review of 2003, and the The Mammals ofAfrica of 2013 edited by T. Butynski and colleagues, place C. doggetti as a subspecies of C. mitis. C. P. Groves in his 2001 Primate Taxonomy classified C. doggetti as a distinct species, which is followed here. Hybridization with C. matis stuhlmanni is reported from the Bwindi Forest and C. ascanius in Gombe Stream National Park, Tanzania. Other hybridizations include C. doggetti x C. denti and C. doggetti x C. kandti. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF82FF80FAEA6DB0FDFDF9CC.taxon	distribution	Distribution. E DR Congo, mountains W of the lakes Edward and Tanganyika (S limit reported at 3 ° 52 ° S, 28 ° 55 " E) extending well W of the main highlands at Nyakanyendje (2 ° 24 ° S, 28 ° 18 ’ E), and into S Uganda, Rwanda, Burundi (E between lakes Kivu and Tanganyika), and NW Tanzania (Bukoba District).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF82FF80FAEA6DB0FDFDF9CC.taxon	description	Descriptive notes. The only available body measurements are from a single adult female head-body 48 - 7 cm, tail 65 - 5 cm (provided by Pocock in 1907). The Silver Monkey is similar to the Blue Monkey (C. mitis), but its pelage is light silvery-brown. Crown, nape, and terminal one-third oftail are black. Tail of the Silver Monkey is often reddish underneath — a distinguishing feature. A buff-speckled frontal band contrasts with a black crown. Body hairs are gray at their base, with five to nine bands of alternating yellow and black; four band pairs occur on the throat.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF82FF80FAEA6DB0FDFDF9CC.taxon	biology_ecology	Habitat. Primary forest, secondary growth forest, and montane forests. The Silver Monkey is also present in edge habitats, although they use mature forests habitats more frequently. B. A. Kaplin reported Silver Monkeys in bamboo forest. They are found at elevations between 770 m in Gombe Stream National Park in Tanzania and 2700 m in Nyungwe Forest Reserve, Rwanda. Rainfall averages 1744 cm / year in areas where they have been studied.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF82FF80FAEA6DB0FDFDF9CC.taxon	food_feeding	Food and Feeding. Silver Monkeys are dietary generalists, but fruit makes up almost 50 % of their diets. Insects make up a considerable proportion of their diets, reported to be 27 % in a study in Rwanda in 1998. They eat young and mature leaves, seeds, flowers, mosses, lichens, gum, and fungi. Insects in their diets include termites, grasshoppers, ants, grubs from rotting wood, and spittlebugs (hemipteran nymphs, Cercopoidea). Dietary diversity increases with decreased fruit availability. Groups living in swamps and riparian forests feed on soft stems of papyrus.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF82FF80FAEA6DB0FDFDF9CC.taxon	breeding	Breeding. There is no specific information available for this species, but breeding is assumed to be similar to the Blue Monkey.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF82FF80FAEA6DB0FDFDF9CC.taxon	activity	Activity patterns. Silver Monkeys are diurnal and arboreal, spending 5 % oftheir time at heights above 20 m in the forest canopy, more than 50 % of their time at 10 - 20 m, 35 % at 5 - 10 m, 3 % between the ground and 5 m, and 2 % on the ground. Locomotion is quadrupedal and includes climbing and leaping.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF82FF80FAEA6DB0FDFDF9CC.taxon	biology_ecology	Movements, Home range and Social organization. Silver Monkeys show fidelity to their home ranges, which were 23 - 28 ha in a study in Nyungwe Forest Reserve, Rwanda. Daily movements are relatively long, like those of Blue Monkeys, at ¢. 1300 m. They live in patriarchal, unimale-multifemale social groups of 20 - 30 individuals. An isolated group in gallery forest in South Ankole grew from 15 to 64 individuals over ten years. Males emigrate from natal groups. Groups may have peripheral males in addition to the dominant male.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF82FF80FAEA6DB0FDFDF9CC.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Least Concern on The [UCN Red List (as C. mitis doggetti). Nevertheless, the Silver Monkey occurs in a region of major habitat loss due to expanding human settlement and agriculture, and it is experiencing a decreasing population trend. Because it is found in several important conservation areas, this decline is not considered to be severe enough to justify its listing in a threatened category.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF85FF80FF2E6368F874FB58.taxon	materials_examined	Virunga Volcanoes, DR Congo. Restricted by C P. Groves in 2001 (following W. C. O. Hill in 1966) to “ near Lake Kivu. ”	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF85FF80FF2E6368F874FB58.taxon	discussion	C. kandti is a member of the matis species group of guenons. J. Kingdon in his 1997 Field Guide to African Mammals, P. Grubb and colleagues in their review of 2003, and The Mammals of Africa of 2013, edited by T. Butynski and colleagues, place C. kandti as a subspecies of C. mitis. Groves in his 2001 Primate Taxonomy classified C. kandti as a distinct species, which is followed here. Hybridization has been reported with C. doggetti in the Mgahinga Gorilla National Park in Uganda. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF85FF80FF2E6368F874FB58.taxon	distribution	Distribution. Endemic to the Albertine Rift, with confirmed presence in remnant Afromontane forests of E DR Congo and SW Uganda (Mt Sabinyo and Mt Mgahinga) in the W Virunga Volcanoes and in W Rwanda (Gishwati and Nyungwe forests); it is also believed to occur at Burunga and Rubengera in the E Virunga Volcanoes.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF85FF80FF2E6368F874FB58.taxon	description	Descriptive notes. Head — body 57 - 65 cm, tail 79 cm; weight 8 - 10 kg (males) and 3.3 - 3.5 kg (females). Body, cheeks, and frontal diadem of the Virungas Golden Monkey are bright golden-yellow, with a contrasting black face, crown, shoulders, limbs, and terminal one-third of tail. Perineum and underside of base of tail are rusty-red. Hairs are yellow at their base, followed by four to seven pairs of alternating red and black bands; yellow bands are broader on flanks, and black bands broader on the back.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF85FF80FF2E6368F874FB58.taxon	biology_ecology	Habitat. Montane forest, with a preference for areas of vegetation with bamboo. Virungas Golden Monkeys also occur in remnant and secondary growth forest. They occur at high elevations of 2100 - 2384 m, with smaller and fewer groups at the higher elevations. Volcanoes National Park, where they were studied by D. Tuyisingize and colleagues in 2006, has an annual rainfall of 1811 mm.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF85FF80FF2E6368F874FB58.taxon	food_feeding	Food and Feeding. Overall dietary diversity of Virungas Golden Monkeysis low, with the top five foods making up 81 - 85 % of their diet. They are primarily folivorous, selecting especially young leaves. They eat fruit, but there is little available where they live. They also eat flowers and bark. Insects made up 16 - 4 % of the diet in a study by D. Twinomugisha, but only 2: 4 % in a study site by Tuyisingize and colleagues in the Volcanoes National Park, Rwanda. Bamboo shoots are important seasonally, up to 27 - 9 % of their time feeding. They raid potato crops and eat bird eggs and soil.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF85FF80FF2E6368F874FB58.taxon	breeding	Breeding. There is no specific information for this species, but Virungas Golden Monkeys are thought to breed throughout the year.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF85FF80FF2E6368F874FB58.taxon	activity	Activity patterns. Virungas Golden Monkeys are diurnal and arboreal. Their locomotion has not been studied specifically, but it is probably similar to other guenons in the mitis species group.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF85FF80FF2E6368F874FB58.taxon	biology_ecology	Movements, Home range and Social organization. Home ranges of Virungas Golden Monkeys are 60 - 74 ha, but one study in Volcanoes National Park in 1984 reported a group of eleven individuals inhabiting a home range of 250 ha. Daily movements are 263 - 898 m. Virungas Golden Monkeys live in male-dominated hierarchical groups. Multimale — multifemale and unimale — multifemale social groups have been reported, with extra males joining groups during the mating season. Group sizes are 3 - 62 individuals, averaging 30 individuals.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF85FF80FF2E6368F874FB58.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Endangered on The IUCN Red List (as C. mitis kandti). The Virungas Golden Monkeyis listed as Class B in the African Convention on the Conservation of Nature and Natural Resources. Its distribution is very restricted, butits precise limits are unknown; the maximum extent of occurrence is estimated at less than 5000 km?. The Virungas Golden Monkey is very dependent on bamboo and its diet overlaps with the Virunga Buffalo (Syncerus mathewsi) and the Mountain Gorilla (Gorilla beringei beringer). Numbers of Virunga Buffalo are increasing in the area, slowing the rate of natural regeneration of the vegetation. Bamboo is illegally harvested despite the protected status of the forests where Virungas Golden Monkeys occur. There is also concern about the unknown effects of climate change on high-elevation forests. The Virungas Golden Monkey occurs in Mgahinga Gorilla National Park in Uganda, Volcanoes National Park in Rwanda, and Virunga and Kahuzi-Biéga national parks in DR Congo.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF85FF81FA2961FCF918F5D2.taxon	vernacular_names	Sykes’s Monkey	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF85FF81FA2961FCF918F5D2.taxon	vernacular_names	French: Cercopithéque a gorge blanche / German: Weiltkehlmeerkatze / Spanish: Cercopiteco de Sykes Other common names: Sykes's Guenon, White-throated Guenon; Kolb's White-collared Monkey (kolbi), Moloney’s White-collared Monkey (moloneyi), Mount Kilimanjaro Sykes’ ' s Monkey (kibonotensis), Patta Island Monkey (phylax), Pousargues’s White-collared Monkey / Tana River Monkey (albotorquatus), Red-eared Sykes’ ' s Monkey (francescae), Samango Monkey (/ abiatus), Schwarz's White-collared Monkey (schwarz), Somalia White-collared Monkey (zammaranoi), Stairs's White-collared Monkey (erythrarchus), Tanzania Sykes's Monkey (monoides), Zanzibar Sykes's Monkey (albogularis)	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF85FF81FA2961FCF918F5D2.taxon	materials_examined	Tanzania, Zanzibar.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF85FF81FA2961FCF918F5D2.taxon	discussion	In his 1997 Field Guide to African Mammals, |]. Kingdon aligned C. nictitans with C. mitis as comprising a “ gentle monkey, ” C. nictitans superspecies group. C. matis in this arrangement included four clusters: Mitis monkeys, white-throated monkeys (including albogularis), silver monkeys (including doggetti and kandti), and blue monkeys — 22 taxa in all, but not pinned down as species or subspecies. This arrangement was followed in The Mammals ofAfrica (Volume 2) edited by T. Butynski and colleagues and published in 2013. The 2003 review of P. Grubb and colleagues likewise adhered to Kingdon’s arrangement. They had two species in the C. nictitans group: C. nictitans (with two subspecies) and C. matus (with all the forms that they considered valid attributed to 16 subspecies, including albogularis, doggetti, and kandti). C. P. Groves in his book 2001 Primate Taxonomy included five species in his mitis species group: C. nictitans with two subspecies (but five are acknowledged here), C. mitis (with seven subspecies), C. doggetti, C. kandi, and C. albogularis (with twelve subspecies: the nominate albogularis, albotorquatus, zammarano, kolbi, kibonotensis, phylax, monoides, moloneyi, francescae, erythrarchus, schwarz, and labiatus); this arrangementis followed here. Grubb and colleagues considered kibonotensis to be a junior synonym of albogularis; phylax and zammaranoi to be junior synonyms of albotorquatus, and schwarzi ajunior synonym of erythrarchus. C. m. stuhlmannii x C. albogularis hybridization is reported in Ngorongoro and Lake Manyara in northern Tanzania. Twelve subspecies recognized.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF85FF81FA2961FCF918F5D2.taxon	distribution	Subspecies and Distribution. C. a. albogularis Sykes, 1831 — Zanzibar I, Tanzania. C. a. albotorquatus de Pousargues, 1896 — N Kenyan coast (Tana River forests). C. a. erythrarchus Peters, 1852 — S Malawi (Mlanje Plateau), E Zimbabwe and most of Mozambique (including the Bazaruto Archipelago). C. a. francescae Thomas, 1902 — N Malawi (Mt Waller and Vipya Plateau, 10 ° 40 ° - 11 ° HO’ 8). C. a. kibonotensis Lonnberg, 1908 — SE Kenya (N to Kilifi Creek) and N Tanzania (inland to Mt Kilimanjaro and Mt Meru). C. a. kolbi Neumann, 1902 — Kenya highlands E of the Rift Valley (Aberdare Mts, Mt Kenya, and Ngong Hills). C. a. labiatus 1. Geoffroy Saint-Hilaire, 1842 — South Africa (Kwazulu-Natal and Eastern Cape provinces); possibly in Lesotho. C. a. moloneyi Sclater, 1894 — S highlands of Tanzania (Poroto Mts, 9 ° S 8.33 ° 40 ’ ' EK) o Udzungwa Mts, Zambia (E of the Luangwa River), and N Malawi (to 9 ° 30 ° S). C. a. monoides 1. Geoffroy Saint-Hilaire, 1841 — E Tanzania, from the seaboard region of Morogoro (6 ° 49 ’ S, 37 ° 40 ’ E) to the Newala District close to the Mozambique border. C. a. phylax Schwarz, 1927 — Pate and Witu Is in the Lamu Archipelago, off the coast of Kenya. C. a. schwarzi Roberts, 1931 — NE South Africa (Limpopo and Mpumalanga provinces) and Swaziland. C. a. zammaranoi de Beaux, 1924 — S Somalia (along theJubba and Webi-Shebelle rivers).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF85FF81FA2961FCF918F5D2.taxon	description	Descriptive notes. Head — body 25 - 68 cm (males) and 20 - 65 cm (females), tail 31 - 95 cm (males) and 26 - 85 cm (females); weight 2: 7 - 11.1 kg (males) and 1.3 - 6 kg (females). Males are larger than females. Sykes’s Monkey is similar to the Blue Monkey (C. matis), but it is generally more uniformly colored, with no scapular band. Body and crown are a relatively light, speckled yellow-olive or ochery-toned with dark limbs. Chin, cheeks, and throat are white. It has long brow fringe projecting forward over eyes and long, pale, or reddish tufts on ears. The “ Zanzibar Sykes’s Monkey ” (C. a. albogularis) is small. Its brow fringe, cheek whiskers, and shoulders are speckled yellowish gray, becoming reddish brown on its dorsum. Its crown is slightly darker than its body. About 60 % of the terminal part oftail is black, with some red hairs on underside of tail base. Its throat patch extends around the neck to make an almost complete collar. Hairs are yellow-gray at their bases, with three pairs of yellow and black bands. “ Pousargues’s White-collared Monkey ” (C. a. albotorquatus) is darker than the nominate subspecies, and its collar is more clearly complete. Its rump is orangey. Underside of tail is red, with red tones on inner thighs. Hairs are yellow-gray at their bases, with pairs of yellow and black bands, as in the nominate subspecies. “ Stairs’s White-collared Monkey ” (C. a. erythrarchus) is somewhat more yellowish. Its posterior dorsum is yellowbrown. Its throat patch is whitish but does not form a collar. It has a caudal patch that sometimes extends completely around tail base. The “ Red-eared Sykes’s Monkey ” (C. a. francescae) is a darker gray, with a short gray collar and red ear tufts; it has reddish hairs on lateral underside of tail and white along the midline. The “ Mount Kiliman-Jaro Sykes’s Monkey ” (C. a. kibonotensis) is very close in description to the nominate subspecies. Ends ofits collar are farther apart at nape, and terminal halfoftail is black. Hairs are gray at their bases, with five pairs of alternating reddish yellow and black bands. “ Kolb’s White-collared Monkey ” (C. a. kolbi) is the largest of the subspecies; it is similar to the Zanzibar Sykes’s Monkey but slightly darker, with a redder dorsum and a nearly complete white collar. Its hairs are yellow-gray at the base, with four pairs of alternating reddish yellow and black bands. The “ Samango Monkey ” (C. a. labiatus) is dark gray, with a slightly yellower posterior dorsum; it has no red buta slightly creamy stripe underits tail base. It has a dark frontal fringe, a partial white collar, and sometimes a white moustache. Hairs are gray at their bases, with three pairs of white and black bands. “ Moloney’s White-collared Monkey ” (C. a. moloneyi) is speckled gray, with light speckled gray whiskers and brow ridge. It has a dark reddish saddle. Crown and nape are dark gray, and throat patch is creamy and extends around the neck as a collar. Distal two-thirds oftail is black, and underside oftail is often reddish. Hairs are gray at their bases, with four pairs of red and black bands. Crown, brow fringe, cheek whiskers, and shoulders of the “ Tanzania Sykes’s Monkey ” (C. a. monoides) are speckled yellowish gray, becoming reddish brown toward its dorsum. About 40 % of the terminal part oftail is black, and there are some red hairs underneath. Its throat patch does not extend to a collar. The “ Patta Island Monkey ” (C. a. phylax) has a more reddish color overall, with a darker crown. Ends of its collar are farther apart at the nape. Tailis black over the terminal half. Hairs are gray at their bases, with five pairs of alternating bands of reddish yellow and black. “ Schwarz’s White-collared Monkey ” (C. a. schwarzi) is less dark and has a less extensive white midline stripe and some red under tail base. The “ Somalia White-collared Monkey ” (C. a. zammaranoi) is an overall dark olive, becoming yellower posteriorly. Its nape is darker than its body and crown. It has no collar and no red underits tail.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF85FF81FA2961FCF918F5D2.taxon	biology_ecology	Habitat. Variable by subspecies, including deciduous, evergreen, riparian forest, and bush woodland (Pousargues’s White-collared Monkey); lowland forest (Pousargues’s White-collared Monkey and the Tanzania Sykes’s Monkey); subtropical, savanna, Acacia (Fabaceae), and swamp forest (Stairs’s White-collared Monkey); coastal forest (Stairs’s White-collared Monkey and the Samango Monkey); montane rainforest (the Tanzania Sykes’s Monkey); and gallery forest (the Somalia White-collared Monkey). Habitats of the Zanzibar Sykes’s Monkey, Moloney’s White-collared Monkey, the Red-eared Sykes’s Monkey, Kolb’s White-collared Monkey, the Mount Kilimanjaro Sykes’s Monkey, the Patta Island Monkey, and Schwarz’s White-collared Monkey have not been explicitly studied, although the Zanzibar Sykes’s Monkey has been reported in coral rag forest, mangrove forest, and edge forest adjacent to plantations. These subspecies are presumed to be similar to other closely related taxa and inhabit tropical forest habitats. The occurrence of Stairs’s White-collared Monkey in swamp and savanna habitats is the result of deforestation on Bazaruto Island, Mozambique, and the restriction of the island subpopulation to the remaining suboptimal habitat.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF85FF81FA2961FCF918F5D2.taxon	food_feeding	Food and Feeding. Few studies of diet and feeding of Sykes’s Monkey have been conducted, but there have been studies, in some detail, on the Zanzibar Sykes’s Monkey, Stairs’s White-collared Monkey, and the Samango Monkey. The Zanzibar Sykes’s Monkey eats fruit and flowers, but it is not clear what proportion ofits diet is composed of these items. It is also reported to raid papaya and banana plantations. Diet of Stairs’s White-collared Monkey was studied in the Cape Vidal region of South Africa, and it contained 43 - 59 % fruit, 22 — 34 % young and mature leaves, 5 - 13 % flowers, ¢. 2: 5 % buds, and 1 - 6 % insects; bark, fungi, and gum were also eaten. Diet and food consumption of the Samango Monkey were studied in Ngoye Forest, South Africa. They were primarily frugivorous with a diet of 84 - 91 % ripe and unripe fruit, 1 - 6 - 3 - 0 % leaves, and 0 - 6 - 2 - 1 % flowers. They also ate bark, gum, fungi, and unspecified animal matter.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF85FF81FA2961FCF918F5D2.taxon	breeding	Breeding. Gestation of the Zanzibar Sykes’s Monkey, Stairs’s White-collared Monkey, and the Samango Monkey is ¢. 5 — 6 months. Menstrual cycles of Stairs’s and Pousargues’s white-collared monkeys are 32 days.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF85FF81FA2961FCF918F5D2.taxon	activity	Activity patterns. Sykes’s Monkeys are diurnal and arboreal. Although their locomotion and postures have not been studied, they are probably similar to other closely related taxa that are predominantly quadrupedal, climbers, and leapers.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF85FF81FA2961FCF918F5D2.taxon	biology_ecology	Movements, Home range and Social organization. Home ranges are 13 - 5 - 15 ha for Stairs’s White-collared Monkey and 14 - 5 ha for Kolb’s White-collared Monkey. Group sizes vary for different subspecies: 15 - 20 individuals for Pousargues’s White-collared Monkey, 30 - 35 for Stairs’s White-collared Monkey, 2 - 34 for the Samango Monkey, and 14 - 18 for Kolb’s White-collared Monkey. The social organization of Stairs’s Whitecollared Monkey is reported to be a hierarchical unimale-multifemale system.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF85FF81FA2961FCF918F5D2.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Least Concern on The [UCN Red List (as C. matis albogularis), including the nominate subspecies albogularis, erythrarchus (as C. m. erythrarchus), kolbi (as C. m. kolbi), moloneyi (as C. m. moloneyi), and monoides (as C. mitis monoides). The subspecies zammaranoi (as C. m. zammaronoi) is classified as Critically Endangered, albotorquatus (as C. m. albotorquatus), labiatus (as C. m. labiatus) as Vulnerable, and francescae (as C. m. francescae) as Data Deficient. The subspecies kibonotensis, phylax, and schwarzi have not been assessed. Although widespread, the Zanzibar Sykes’s Monkey is declining where it occurs outside protected areas. Pousargues’s White-collared Monkey has an extent of occurrence less than 20,000 km?, and it is experiencing population declines because of habitat fragmentation and habitat loss and degradation from agriculture. Stairs’s White-collared Monkey has a wide distribution and is locally common, although it is experiencing population declines in some areas. The Red-eared Sykes’s Monkey has no current information on its threats and population status. Much of the distribution of Kolb’s White-collared Monkey is within protected areas, including Aberdare National Park, Mount Kenya National Park, and several forest reserves. It is locally common and relatively well protected. The Samango Monkey is endemic to South Africa in highly fragmented, discrete, and isolated forests. Its past decline exceeds 30 % over 27 years (three generations). Habitat loss is widespread in its already fragmented habitat due to past logging practices and current forestry plantations. Moloney’s White-collared Monkey is widespread in suitable habitat and occurs in several protected areas. Although it experiences population declines, they are not severe enough to warrant a threatened status. The Tanzania Sykes’s Monkey is also widespread in suitable habitat, even though it is probably declining in all areas outside of protected areas because of habitat loss and degradation. The population of the Somalia White-collared Monkey has been estimated at only 200 - 500 individuals, and it is declining, with no more than 50 mature individuals in any of the subpopulations.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF86FF83FF236740FA0AF442.taxon	materials_examined	Angola, 300 miles (483 km) inland from Bembe.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF86FF83FF236740FA0AF442.taxon	discussion	In 2001, C. P. Groves recognized all of the named subspecies listed here except for sharper, which he believed to be a junior synonym of palliatus. C. sharpei was considered a valid subspecies of C. angolensis by U. Rahm in 1970, J. Kingdon in his 1971 review of East African primates, and P. Napier in 1985. In 1981, W. Rodgers provided information on the distributions of palliatus and sharpei that showed they were not isolated as they were thought to be and suggested they were not distinct taxa. Nevertheless, a molecular genetic study of palliatus across its distribution by M. McDonald and H. Hamilton in 2010 demonstrated distinct haplotypes in Kenyan and Tanzanian populations, which, combined with pelage differences highlighted by Rahm, Napier, and Kingdon, supported the validity of sharpei in central and southern Tanzania as a subspecies distinct from palliatus in Kenya. The form prigogine: could be synonymous with cordieri, but it is recognized as a subspecies of C. angolensis here. An unnamed subspecies of C. angolensis, the “ Nkungwe’s Angolan Colobus ” or “ Mahale Mountains Angola Colobus ” was listed by Groves in 2001 and P. Grubb and colleagues in their taxonomic review in 2003. It is an isolated population reported by T. Nishida and colleagues in 1981 from Mount Nkwunge and ridges of the northern Mahale Mountains as far as Mount Pasagula to the north and Mount Kahoko and Mount Sibindi to the south. It wasillustrated and described in a field guide of primates of the Mahale Mountains by T. Butynski and Y. de Jong in 2009, but it remains without a formal scientific trinomial. Seven subspecies recognized.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF86FF83FF236740FA0AF442.taxon	distribution	Subspecies and Distribution. C. a. angolensisSclater, 1860 — DRCongo (SofthegreatbendoftheCongoRiver), NEAngola, andNWZambia, Stoc. 12 ° SandWto ¢. 16 ° E. C. a. cordier: Rahm, 1959 — EDRCongoWoftheLualabaRiver, betweentheLowaandLuamariverstoclosetoLakeKivuintheEat ¢. 28 ° E (whereithybridizeswiththesubspeciesruwenzorii), andtothehighlandsWofLakeTanganyika, whereitextendstoc. 6 ° S. C. a. cotton: Lydekker, 1905 — NEDRCongoEoftheCongo-L. ualabariversystem, fromtheNbankoftheCongoRiver (c. 20 ° E) NtotheUeleRiver, EtoLakeAlbert, andStotheLindiRiver; formsahybridzonewiththesubspeciesruwenzoriiintheSofitsdistribution. C. a. palliatusPeters, 1868 — SEKenyaandNETanzania. C. a. prigogineiVerheyen, 1959 — EDRCongo (MtKabobo). C. a. ruwenzoriiThomas, 1901 — SEDRCongo, SEUganda, WRwanda, WBurundi, andNWTanzania; alongbothsidesoftheWRift, fromtheSemlikiValley, theRuwenzoriMts, andMtKahuzitotheshoresofLakeTanganyika. C. a. sharpei Thomas, 1902 — C & S Tanzania (as far W as Lake Rukwa), N Malawi, NW Zambia, and possibly occurring in N Mozambique.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF86FF83FF236740FA0AF442.taxon	description	Descriptive notes. Head — body 55 - 66 cm (males) and 48 - 59 cm (females), tail 76 92 cm (males) and 63 - 76 cm (females); weight 7.6 - 12.6 kg (males) and 6.4 - 9.2 kg (females). The Angolan Colobus is generally black and characterized by long white epaulettes on shoulders and fluffy white circumfacial hair, the latter forming laterally elongated tufts on cheeks that resolve into a crescent shape. Tail may be tufted in males (not tufted in females), with varying extents of white (sometimes entirely white or with white restricted to the tuft). Two subspecies groups are recognized. The first is characterized by large epaulettes and a white pubic band (“ Sclater’s Angolan Colobus, ” C. a. angolensis; “ Peters’s Angolan Colobus, ” C. a. palliatus; “ Adolf Friedrich’s Angolan Colobus, ” C. a. ruwenzorii; and “ Sharpe’s Angolan Colobus, ” C. a. sharper). The other subspecies group has thin epaulettes and no white band on the pubic region (“ Cordier’s Angolan Colobus, ” C. a. cordieri; “ Powell-Cotton’s Angolan Colobus, ” C. a. cottons; and “ Prigogine’s Angolan Colobus, ” C. a. prigoginei). These two subspecies groups are linked by the unnamed Nkungwe Angolan Colobus that has well-developed epaulettes but lacks forehead and pubic bands. It is possible that some or all described subspecies may be distinct species. Sharpe’s Angolan Colobus is notable for its particularly long epaulettes, extending to its forearms, and bushy white side-whiskers. Prigogine’s Angolan Colobus has the smallest epaulettes and relatively reduced white side-whiskers.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF86FF83FF236740FA0AF442.taxon	biology_ecology	Habitat. A variety of forest types, from primary and secondary lowland rainforest and gallery forest to montane forest up to elevations of 3000 m and as high as 4000 m on Mount Ruwenzori. Although Angolan Colobus prefer the very highest levels of the forest canopy, including emergent trees, individuals will occasionally descend to the ground near streams to feed upon herbaceous vegetation.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF86FF83FF236740FA0AF442.taxon	food_feeding	Food and Feeding. Diets of Angolan Colobus are mainly young leaves, along with shoots, seeds, unripe fruit, mature leaves, flowers, lichen, and termite clay. Water is obtained primarily from their food.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF86FF83FF236740FA0AF442.taxon	breeding	Breeding. Female Angolan Colobus do not display a sexual swelling during the periovulatory period, and there is no perineal organ in the male. There is a definite birth season (May-December) in at least part of its distribution. A single young is born. The Natal coat is white, gradually changing through gray to black during the first six months.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF86FF83FF236740FA0AF442.taxon	activity	Activity patterns. The Angolan Colobus is diurnal and arboreal. At Nyungwe in Rwanda, its activity budget is 42 % feeding, 32 % resting, 20 % moving, 5 % engaging in social behavior, and 1 % other activities. This population has unusually large group sizes (over 300 individuals), which may explain unusually high amounts of time spent feeding and moving and a relatively small amount of time devoted to resting, especially for a colobine. In a study in the Ituri forest of the DR Congo, groups of Angolan Colobus rested 43 % of their time, fed 27 %, moved 24 %, engaged in social behavior 5 %, and showed other behaviors 1 %.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF86FF83FF236740FA0AF442.taxon	biology_ecology	Movements, Home range and Social organization. Daily movement has been reported at 983 m, with a home range of 371 ha. The Angolan Colobus generally lives in small family groups of 3 — 4 individuals, with a single male present. These family groups often form “ troops ” of up to 25 - 30 individuals, in which more than one male is present. One population in Rwanda (Nyungwe Forest) is exceptional in having a troop of more than 300 individuals, using a 2440 ha home range. Little is known about the dispersal patterns of Angolan Colobus.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF86FF83FF236740FA0AF442.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Least Concern on The IUCN Red List. The subspecies prigoginei is classified as Endangered, ruwenzorii as Vulnerable, cordieri as Near Threatened, and angolensis, cottoni, palliatus, and sharpei (under C. a. palliatus) as Least Concern. The undescribed Nkungwe taxon has not been assessed, but it is believed to be threatened. The Angolan Colobusis listed as Class B in the African Convention on the Conservation of Nature and Natural Resources. Although threatened in some parts ofits distribution (particularly in East Africa and the Albertine Rift), it remains widespread and relatively common in suitable habitat. Deforestation for timber and conversion of land for agricultural use are major threats, and isolated populations such as those in the Eastern Arc and coastal forests are especially vulnerable. It is also hunted for meat, especially in the Congo Basin. Populations in the eastern part ofits distribution are much more fragmented than those across the Congo Basin. The wide-ranging nominate form (Sclater’s Angolan Colobus) is known to occur in Salonga National Park in DR Congo and Mweru Wantipa National Park in Zambia. Peters’s Angolan Colobus occurs in isolated forests in south-eastern Kenya, restricted to the southern coastal forests of the Kwale District; a comprehensive survey located 55 populations. Although locally threatened from habitat loss and hunting in some parts of its highly fragmented distribution, Peters’s Angolan Colobus remains widespread and relatively common. The total population is estimated at 3100 - 5000 individuals (560 - 900 groups), with the population in Shimba Hills Forest Reserve being the largest at ¢. 2400 individuals (c. 15 ind / km?). Sharpe’s Angolan Colobus is likewise discontinuously distributed in forest patches in the Southern Highlands and coastal and gallery forests of southern and eastern Tanzania. It occurs in Udzungwa Mountains National Park and Gendagenda South Forest Reserve. Cordier’s Angolan Colobus has declined as a result of increased hunting. It is not known whether it occurs in any protected areas. Powell-Cotton’s Angolan Colobus is locally threatened from habitat loss and hunting in some parts ofits distribution but remains widespread. It occurs in Kahuzi-Biéga National Park, Virunga National Park, and Tayna Gorilla Reserve in DR Congo. Prigogine’s Angolan Colobus is confined entirely to Mount Kabobo in eastern DR Congo, where it is at continuing risk of habitat loss from agricultural expansion and hunting. It is not known whether it occurs in any protected areas. Adolf Friedrich’s Angolan Colobus is restricted to a few sites in the Albertine Rift region, where its habitat is fragmented and at risk from continued degradation. Hunting is also a threat. It occurs in Ruwenzori National Park in Uganda, Nyungwe Forest National Park in Rwanda, Kaya Gonja Nature Reserve in Kenya, and Mahale Mountains National Park in Tanzania.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF86FF8DFA2C6916F836FE4A.taxon	materials_examined	Ethiopia, Gojjam and Kulla.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF86FF8DFA2C6916F836FE4A.taxon	discussion	The more thinly furred lowland population of C. guereza along the Omo River perhaps should be separated as a different subspecies poliurus named by Thomas in 1901. Further taxonomic work is needed to assess the validity of some of these taxa and identify the possible existence of previously undescribed taxa. Eight subspecies recognized.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF86FF8DFA2C6916F836FE4A.taxon	distribution	Subspecies and Distribution. C. g. guerezaRuppell, 1835 — WEthiopia, intheforestedareasoftheEthiopianHighlandsWoftheRiftValley, extendingintolowlandforestsalongtheOmoRivertotheSandtheBlueNiletotheW. C. g. caudatusThomas, 1885 — SKenyaandNTanzania, inthemontaneforestsofMtKilimanjaroandMtMeru, aswellastheadjoiningforestsatslightlylowerelevations (Kahé, MomelaLakesinArushaNationalPark). C. g. dodingaeMatschie, 1913 — SESudan, knownonlyfromtheDidingaHills. C. g. gallarumNeumann, 1902 — ECEthiopia, intheEthiopianHighlandsEoftheRiftValley; theprecisedistributionalboundariesareunclear, andthoseoftheBaleMtsmaybedistinctfromthoseoftheArsi (= Arussi) Mts. C. g. kikuyuensisLonnberg, 1912 — CKenya, intheKenyanHighlands, EoftheRiftValley, fromtheNgongEscarpmenttoMtKenyaandtheAberdareRange. C. g. matschietNeumann, 1899 — SWUganda (MtElgon), WKenya (WoftheRiftValley, NtoMtElgon, andinsomeforestswithintheRiftitself, asnearLakeNaivasha), andNWTanzania (NofNgorongoroCraterandtheGrumetiRiver). C. g. occidentalisRochebrune, 1887 — SCameroon (WtoBatouriandLornie, whereitisapparentlysympatricwiththeBlackColobus (C. satanas), andNacrosstheSanagaRivertoYabassi), SWCentralAfricanRepublic, Chad, SWSudan, NDRCongo (StotheCongoRiverandtoc. 1 ° 25 ” SintheIturiForest), NEGabon (MakokouandBelingadistricts), NRepublicoftheCongo, WUganda (WoftheNile), andNWRwanda; possiblyinEquatorialGuineaandtheDongaRiverregionofSENigeria. C. g. percivali Heller, 1913 — WC Kenya, restricted to a small area of forest around Mt Gargues in the Matthews Range.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF86FF8DFA2C6916F836FE4A.taxon	description	Descriptive notes. Head-body 53 - 75 cm (males) and 49.5 - 67.3 cm (females), tail 52 - 90 cm (males) and 50 - 80 cm (females); weight 8 - 13.5 kg (males, exceptionally up to 23 kg) and 5.5 - 10.2 kg (females). The Guereza is the largest of the African colobines. It is a somewhat variable species, but it always has short black hair with long plumes of white on the tail and running down the sides from shoulders to haunch, the latter curving across the lumbar region to form a U-shaped mantle or flank veil. Exact arrangement and length of white fur depends greatly on locality, although, as a rule, individuals native to higher elevations have longer coats than do those from lowland forests. Tail is usually longer than head-body length and sports a long, full white tuft that is either restricted to the terminal zone or occupies nearly the entire length. There is a white thigh stripe, and short but bushy white cheek hairs make up part of a whitish fringe, which includes a bushy beard, encircling the hairless, darkgray face. Nose is hooked and has a tendency to overhang the mouth somewhat. Sexes are distinguished by size and fur pattern in the perineal region: males have a white semi-circle, whereas in females this pattern is bisected by a black line. Various morphs are also known in addition to the recognized subspecies. Brown and white Guerezas have been seen in parts of Ethiopia, and there is even a small population of aberrant, entirely white individuals living near Nanyuki on Mount Kenya. In general, subspecies may be divided into two groups: (1) an eastern group (the “ Mount Kilimanjaro Guereza, ” C. g. caudatus; the “ Mount Kenya Guereza, ” C. g. kikuyuensis; and the “ Mount Uaraguess Guereza, ” C. g. percivali), in which the tail is not usually longer than the head-body length and is largely (more than 70 %) occupied by the bushy white tuft, and the white mantleis very long, extending well up onto the dorsum; and (2) a northern and western group (the “ Omo River Guereza, ” C. g. guereza; the “ Djaffa Mountains Guereza, ” C. g. gallarum; the “ Mau Forest Guereza, ” C. g. matschiei; the “ Dodinga Hills Guereza, ” C. g. dodingae, and the “ Western Guereza, ” C. g. occidentalis), in which the tail is longer than the head-body length and half or less is occupied by the white tuft, and the mantle is shorter and thinner.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF86FF8DFA2C6916F836FE4A.taxon	biology_ecology	Habitat. Primary and secondary lowland deciduous and evergreen forest, and medium and montane tropical moist forest, gallery forest, swamp forest, and wooded grassland, especially those near rivers and lakes. Elevational range is up to 3300 m. The Guereza is often found in disturbed, secondary, and successional forests, and it prefers degraded forests to old growth when both are available. This preferenceis likely attributable to high species diversity of food trees in some secondary growth forests, and it may also be explicable in terms of milder chemical defenses in leaves of secondary growth species. The Guereza is also frequently found in areas under human use, such as eucalyptus plantations, which are perhaps visited to make up for nutritional deficiencies. It prefers the upper canopy of the forest.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF86FF8DFA2C6916F836FE4A.taxon	food_feeding	Food and Feeding. The Guereza eats mainly leaves and fruit. The diet is quite variable, however, as would be expected in a species with such a wide range of distribution and habitat types. Proportions of these types of food relative to one another varies by study site and time of year, often with leaves making up more than half to most of the diet, but with fruit sometimes predominating. Fleshy fruits are usually eaten when unripe, with consumption being reduced as they fully ripen, likely to avoid competition with other primate species that prefer ripe fruit. While a number of species of plants are exploited, often only a few make up the majority of the diet at a specific site. The usual feeding pattern of Guerezas is to select young leaves, but in times of scarcity, to rely on mature leaves and fruit. Their use of mature leaves can vary widely, however, across forests, and even between groups in the same forest. Other foods eaten include bark and wood, seeds, flowers, petioles, lianas, and arthropods. They occasionally go to the ground to feed on termite clay, aquatic plants such as duckweed, soil, and even concrete from buildings.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF86FF8DFA2C6916F836FE4A.taxon	breeding	Breeding. Reproductive cycle of the Guereza lasts c. 24 days, with females being receptive from c. 5 days before ovulation until 2 - 3 days after. Females do not display a sexual swelling during the periovulatory period, and there is no perineal organ in the male. The interbirth interval is ¢. 22 months, with a minimum of 16 months. Birth rates are higher in habitats that are more forested. Copulation is initiated roughly equally between males and females. Solicitation behavior includes approaching a prospective partner and performing low-intensity mouth clicking or tongue-smacking. To present, a female will stand with her hindquarters facing a male and with her forearms bent, holding her body low without raising her tail. The copulation posture consists of the male grasping the female’s ankles and trunk from behind. Copulations usually occur among group members, but extragroup copulations have been observed in some populations. In multimale groups, females will sometimes mate with more than one male. The gestation period in the wild is ¢. 158 days (5 - 2 months), but it has been estimated to be 170 days (5 - 6 months) in captivity. A single young is born; births occur throughout the year, but with certain seasonal peaks. The neonate has white fur with pink to red skin, in stark contrast with the black-and-white adults. By three weeks old, the face and ears start to darken and become gray. The natal coat and skin continue to blacken, reaching adult coloration ¢. 3 — 4 months of age, with males attaining adult coloration sooner than females. Some gray coloration can persist above the ears on top of the head through 6 - 7 months after birth. In the wild, infant mortality in a small sample exceeded one-third of infants dying before their pelage changed, but mortality was lower after the color change. High infant mortality is also observed elsewhere in the wild. Infants are always carried by the mother ventrally, with the infant grasping its mother’s fur during group movements. In the wild and captivity, infants are the focus of attention of other members of the group, especially females, and are often handled or carried by them, even shortly after birth and with the mothers’ full tolerance. Infants are often uncomfortable, however, with being held by individuals other than their mothers. Males are generally disinterested in infants at birth, but their interest increases after infants are 4 - 5 weeks old. Infantcide has been observed, usually committed by non-group or newly immigrated males. Sexual maturity is reached at c. 4 years in females and 4 - 6 years in males. Guerezas can live over 30 years in captivity.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF86FF8DFA2C6916F836FE4A.taxon	activity	Activity patterns. The Guereza is diurnal and arboreal. Across locations in East Central Africa, its activity budget averages 52 - 63 % resting, 19 - 26 % feeding, 2 - 22 % moving, 5 - 11 % engaging in social behavior, and 1 - 7 % miscellaneous other activities. More than half the day is spent resting. The next most common activity is feeding, which also takes up a significant amount of time, and feeding bouts are followed by rest periods in which gases generated by gastric fermentation are quietly burped out. Other activities that occurfar less frequently include grooming and play. While a group is resting, the male often rests higher and away from the group, possibly to detect other groups and raptors. During group movements, various members stay near one another and follow a leading individual, often in single file and along a similar path. Guerezas leave the area around their sleeping trees one to several hours after sunrise and retire to sleeping trees by sunset. The adult male utters a series of loud, croaking roars, sometimes evidently as a means of intergroup spacing (answered by roaring from males of other groups), sometimes stimulated by disturbances such a dog barking or thunder, and sometimes for no detectable reason. The roar series is generally accompanied by noisy jumping and dropping through the canopy (jumping-roaring display). All species of Colobus make these roars and associated displays, but they differ slightly from one species to another.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF86FF8DFA2C6916F836FE4A.taxon	biology_ecology	Movements, Home range and Social organization. In long-term studies, single-group daily movements average 252 - 734 m, ranging from aslittle as 62 m / day to more than 1360 m / day. Home rangesize is variable by study location, with estimates ranging from just over 7 - 5 ha to 100 ha; most estimates are usually ¢. 20 ha or less. Home ranges can overlap. There are also core areas within home ranges that are significantly smaller than the overall home range. Guerezas normally live in small, cohesive groups of 6 - 10 individuals, although groups of up to 40 have been reported in the southern gallery forests of the Central African Republic. Group size tends to be larger in contiguous than fragmented or riparian forest, and it increases with elevation. Groups generally consist of a single adult male (although several can be present in larger groups), several adult females, and immature individuals. Multimale groups are unstable and often relapse into a single-male structure. Guereza groups appear to be matrilineally organized, with females and immature individuals forming the closely bonded core of the group. Adult females in groups are often closely related, but this is not true for all females. Close adult female relatives may also occur among neighboring groups. Even though females are typically philopatric, they may disperse when groups dissolve. In captivity, females do not exhibit a hierarchy of rank, but in the wild, some adult females have dominance relationships. Males typically disperse as subadults or adults. In Kibale Forest National Park, Uganda, close adult male relatives sometimes occur among neighboring groups, but overall low levels of among-group-adult-male relatedness suggest that males often disperse beyond their neighbors. They may become solitary or join bachelor groups, they may immigrate into other groups by joining on the periphery or staging takeovers, and they may also disperse secondarily. Males that take over groups often have tenure lengths reaching five years or more. Multimale groups can consist of father-son pairs, but some apparently contain unrelated males. In mult-male groups, one male is typically dominant over the others and interactions between adult males are aggressive, with some males eventually being forced out. Adult males rarely groom others in the group.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF86FF8DFA2C6916F836FE4A.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Least Concern on The [UCN Red List. The subspecies percivaliis classified as Endangered, guereza, caudatus, kikuyuensis, and occidentalis are classified as Least Concern, and dodingae, gallarum, and matschiei as Data Deficient. The Guereza is listed as Class B in the African Convention on the Conservation of Nature and Natural Resources. It is among the least threatened of the colobines due to its propensity to travel on the ground (able to travel between different forest fragments) and to use dry and gallery forest. Although very widespread and still locally abundant in many areas, Guerezas are nevertheless threatened in parts of their distribution by habitat loss through deforestation for timber and conversion to exotic forest plantations and agricultural land. They are one of the few primates that are generally considered able to cope with habitat degradation. Overuse of the forest and intense disturbance can, however, lead to a decline in their numbers. Clearing of forest for firewood contributes to population declines, but commercial logging poses a larger threat. Commercial uses include wood clearing for brewing beer, fuel, gin distillation, and charcoal production. Even if a forest is not cleared, there are also a number of ways Guereza habitat can be degraded: e. g. cutting of trees for timber, charcoal, firewood, tools and other uses; gold mining; livestock traffic; road building; agriculture; vandalism; poaching; extraction of plant foods; and stripping of bark from trees for medicine. Guerezas sometime respond positively to logging, and they are sometimes found in higher densities in logged areas than unlogged, possibly because of an increase in preferred food trees. In western Uganda, clearing of forest fragments has caused a reduction in numbers of Guerezas by more than 50 % over an 8 year period. Conifer plantations are unsuitable for Guerezas. In recent years, the African cherry (Prunus africana, Rosacae), a sometimes favored food of Guerezas, has declined across sub-Saharan Africa, which has resulted in a corresponding decline in the monkeys that rely on it. In some places, human activities have increased concentrations of African Savanna Elephants (Loxodonta africana), as in Murchison Falls National Park, Uganda, resulting in the destruction of habitat used by Guerezas. Hunting can also be a threat, butitis variable in its occurrence throughout their distribution. In Gabon, for example, hunted populations have declined by 88 %. Guerezas are sold as bushmeat in northern DR Congo, but in Uganda and elsewhere in East Africa they are generally not hunted. Crop-raiding by Guerezas does occur, albeit at low levels. The Guereza has been hunted commercially for centuries for its beautiful pelt (Marco Polo is said to have found colobus capes during his journey to China). During the late 19 " century, they were hunted almost to the point of extinction in some areas; the trade reached its zenith just prior to World War I when “ monkey fur ” was at the height of fashion in Europe. It is estimated that from one to two million colobus were killed during the peak of demand. Skins themselves were used either for trimming coats or made into rugs or wall-hangings. Thankfully, the fad eventually fell out of favor, and Guerezas have since made an almost complete recovery in numbers. A limited amount ofillegal hunting continues to the present; however, most of this is by African tribesmen who use Guerezas for food and in traditional ornamentation. In some areas, souvenirs made from Guereza fur are still sold to tourists. Firearms and traditional weapons are used for hunting. The Omo River Guereza is at most only threatened in parts of its distribution; it is found in Simien Mountains, Awash, and Omo national parks in Ethiopia. The Mount Uaraguess Guereza is at risk from hunting for its skin. It is found in the Matthews Range Forest Reserve in Kenya. The Dodinga Hills Guereza is restricted to the Dodinga Mountains in southern Sudan, where the last known record was from the 1960 s. There is no current information on the status of this subspecies, largely because of inadequate surveys, and it may well be at risk of extinction. It is not found in any protected area. The Western Guereza is perhaps locally threatened in parts of its distribution, but remains widespread. It is found in Dja Biosphere Reserve in Cameroon; Manovo-Gounda-Saint Floris National Park in the Central African Republic; Odzala and Nouabalé-Ndoki national parks in the Republic of the Congo; Garamba and Virunga national parks in DR Congo; and Bwindi Impenetrable, Murchison Falls, Kibale Forest, Mgahinga Gorilla, and Ruwenzori national parks, Budongo, Bugoma, Kalinzu, Kasyoha-Kitomi, Itwara, Mount Kadam forest reserves, and Toro Game Reserve in Uganda. It may also be found in Gashaka-Gumti National Park and Ngelnyaki Forest Reserve in Nigeria. Although the Mount Kilimanjaro Guereza has a relatively confined distribution, there are no obvious major threats. It is found in Arusha and Kilimanjaro national parks in Tanzania. The Djaffa Mountains Guereza may be declining in known parts ofits distribution because of habitat loss from agriculture. Further survey work is needed to better establishits distributional limits and population status. It is found in Bale Mountains National Park and Menagasha National Forest in Ethiopia. The Mount Kenya Guereza is very abundant within its limited distribution, which is largely protected (Aberdare and Mount Kenya national parks in Kenya), and there are no obvious major threats at present. The Mau Forest Guereza may be declining within its limited distribution because of habitat loss to agriculture, although little is known about its status and habitat requirements and further surveys are required. It is found in at least six protected areas: Lake Naivasha, Lake Nakuru, and Mount Elgon national parks and Masai-Mara Game Reserve in Kenya; Mount Elgon National Park in Uganda; and Serengeti National Park in Tanzania.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF88FF8DFA2C64EEF6B7F336.taxon	materials_examined	Sierra Leone (probably Sherbro Island).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF88FF8DFA2C64EEF6B7F336.taxon	discussion	Black-and-white colobus monkeys with uncertain taxonomic status — a hybrid swarm (previously referred to as “ Doll man’s Colobus ” dollmani) of C. polykomos x C. vellerosus exists between the Sassandra River and the Bandama River (and its eastern tributary, the Nzi) in Ivory Coast. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF88FF8DFA2C64EEF6B7F336.taxon	distribution	Distribution. Guinea Bissau, Guinea, Sierra Leone, Liberia, and SW Ivory Coast (E to the Sassandra River); possibly in S Senegal and Gambia.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF88FF8DFA2C64EEF6B7F336.taxon	description	Descriptive notes. Head-body 58 - 68 cm (males) and 55 - 65 cm (females), tail 72 — 100 cm (males) and 81 - 96 cm (females); weight 6: 6 — 10 kg (males) and 6.7 - 8.3 kg (females). The King Colobusis a robust, long-tailed colobine, with its tail being more than 160 % of head-body length. Body, crown, and limbs are mostly black. Crown has no whorl, with its front and sides of head and neck having long gray-white hairs that gradually become intermingled with black on sides of the neck and shoulders (where hairs form long epaulettes). White hairs gradually disappear on flanks and innersides of humeri. Tail is white and untufted. Facial skin is black, with a prominent nose that often protrudes beyond the mouth. Ischial callosities are margined with white fur ventrally in both sexes, with the white extending up between callosities and down toward genitalia. Cranium has low, rounded orbits and a weakly developed rounded supraorbital torus with no concavity at the glabella. Pyriform aperture is low and narrow.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF88FF8DFA2C64EEF6B7F336.taxon	biology_ecology	Habitat. A variety of primary and secondary forest habitats, including dry semi-deciduous, moist evergreen, and riparian forests. The King Colobus is rarely found in degraded habitat, but it does sometimes occur in secondary forests. It is often seen in tall emergenttrees.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF88FF8DFA2C64EEF6B7F336.taxon	food_feeding	Food and Feeding. The King Colobus eats mainly young leaves, mature leaves, and unripe seeds, supplemented with shoots, fruits (mainly unripe), flowers, and termite clay. It relies heavily on lianas and leguminous plants. Water is obtained mainly from their food, and when necessary, individuals can live for up to several months on little but mature leaves.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF88FF8DFA2C64EEF6B7F336.taxon	breeding	Breeding. Female King Colobus do not display a sexual swelling when receptive, and there is no perineal organ in the male. Females initiate copulations. One offspring is born in December — February after a gestation of 170 days. The skin of the infants is pink for the first few months oflife and covered with a short, all-white natal coat. Every mature female in the group takes part in the rearing of the young. One individual lived for 27 years in the Erfurt Zoo, Germany.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF88FF8DFA2C64EEF6B7F336.taxon	activity	Activity patterns. King Colobus are diurnal and arboreal. At Tiwai Island Wildlife Sanctuary, G. Dasilva found them to spend 61 % of their time resting, 28 % feeding, 9 % moving, 1 % engaging in social behavior, and 1 % miscellaneous other activities. Low levels of activity might be a behavioral strategy to conserve energy, particularly when food is scarce. The King Colobus adopts a hunched posture during rainstorms and periods of low temperature, which is likely related to thermoregulation.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF88FF8DFA2C64EEF6B7F336.taxon	biology_ecology	Movements, Home range and Social organization. Daily movements of King Colobus average 637 - 830 m, with home ranges of 24 - 83 ha. Group size is typically 9 - 19 individuals, with 1 - 3 adult males and 4 - 6 adult females, although smaller groups of 3 - 6 individuals have been reported. Males disperse from their natal group, but there is also evidence of female movement. Females typically form bonds in a group, while adult males form dominance hierarchies and rarely interact.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF88FF8DFA2C64EEF6B7F336.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Vulnerable on The IUCN Red List. The King Colobusis listed as Class A in the African Convention on the Conservation of Nature and Natural Resources. Although its total distribution is still extensive, habitat loss and hunting for its meat and pelt are becoming increasing threats and are fragmenting remaining populations. It does not survive well in degraded areas and requires some primary forest to thrive. The King Colobus occurs in nine protected areas, but protection from hunting is still problematic in some of them: Tai Forest National Park in Ivory Coast; Sapo National Park in Liberia; Tiwai Island Wildlife Sanctuary, Gola Forest Reserve, and Outamba-Kilimi National Park in Sierra Leone; and Haut Niger National Park, Mount Nimba Strict Nature Reserve, and Massif du Ziama and Diecke biosphere reserves in Guinea.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF88FF8EFA286806F9D2FDA8.taxon	materials_examined	Equatorial Guinea, Fernando Poo (= Bioko Island).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF88FF8EFA286806F9D2FDA8.taxon	discussion	There is very little external difference between the two subspecies of C. satanas. Two subspecies recognized.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF88FF8EFA286806F9D2FDA8.taxon	distribution	Subspecies and Distribution. C. s. satanasWaterhouse, 1838 — Bioko 1 (EquatorialGuinea). C. s. anthracinus Le Conte, 1857 — E & SW Cameroon (S of Sanaga River and as far E as the Batouri and Lomié districts), Equatorial Guinea, W & C Gabon (inland to Lopé National Park and with unconfirmed reports of sparse populations E of the Ogooué and Ivindo rivers), and W Republic of the Congo; possibly in NW Angola (Cabinda).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF88FF8EFA286806F9D2FDA8.taxon	description	Descriptive notes. Head-body 50 - 77 cm, tail ¢. 80 cm; weight 10 - 15 kg (males) and 10 - 11 kg (females). The Black Colobusis a large, entirely black colobine, with an untufted tail. There is a pronounced mantle, and hairs on cheeks and forehead are quite long. An occipital crest of hair rises from a whorl on the front of the crown, and there is a prominent brow fringe. Face and ears are naked; nose is comparatively short. Skull is less prognathous than in other species of Colobus, and the braincase has a distinctive transverse “ saddle ” behind the bregma. The “ Bioko Black Colobus ” (C. s. satanas) has longer hair than the mainland form, the “ Gabon Black Colobus ™ (C. s. anthracinus). The roaring call of the adult male is primitive; it is preceded by a loud “ sneeze ” and is quite different from those of other Colobus.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF88FF8EFA286806F9D2FDA8.taxon	biology_ecology	Habitat. Dense primary rainforest, mature secondary rainforest, and coastal evergreen, lowland moist, montane, and swamp forest; occasionally coastal sand dune and wooded meadows. The elevational range of the Black Colobus is 500 - 3011 m. It appears unable to survive in degraded secondary forest and is rare or absent in forests where logging has reduced canopy height. On Bioko Island, it has been observed in montane heathland. It avoids areas disturbed by humans. It is typically found high in the canopy.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF88FF8EFA286806F9D2FDA8.taxon	food_feeding	Food and Feeding. Black Colobus eat large quantities of seeds, along with young leaves and mature leaves (particularly those of lianas), unripe fruits, flowers, and buds. Mature leaves are less favored, and they are only eaten with regularity in sand-dune forest. Sodium-rich soil is also consumed. The Black Colobus has a remarkable tolerance for high concentrations of poisonous tannins in leaves and, as a consequence, is able to live in areas where other colobines cannot. It also appears to rely more heavily upon seeds as a dietary component. When food is abundant, groups may feed in the same tree for several days.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF88FF8EFA286806F9D2FDA8.taxon	breeding	Breeding. During the periovulatory period of the Black Colobus, the clitoris protrudes, but females do not otherwise display a sexual swelling. There is no perineal organ in the male. A single young is born after a gestation of 195 days. Infants are brown, not white as in other species of Colobus.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF88FF8EFA286806F9D2FDA8.taxon	activity	Activity patterns. Black Colobus are diurnal and arboreal. At Douala-Edea in Cameroon, D. McKey and P. Waterman found them to spend 60 % of their time resting, 23 % feeding, 13 % engaging in social behavior, and 4 % moving.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF88FF8EFA286806F9D2FDA8.taxon	biology_ecology	Movements, Home range and Social organization. Black Colobus move up to 850 m / day in home ranges of 70 - 570 ha. Their groups contain 7 - 30 individuals, but on Bioko Island, they are typically smaller (7 - 10 individuals). Groups usually contain several adult males. In 1978, the population density in Foret des Abeilles in central Gabon was estimated at ¢. 25 - 30 ind / km?, and in the Douala-Edéa Wildlife Reserve in Cameroon at ¢. 38 ind / km?.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF88FF8EFA286806F9D2FDA8.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Vulnerable on The IUCN Red List, including the subspecies anthracinus, but the nominate subspecies satanas is classified as Endangered. The Black Colobusis listed as Class B in the African Convention on the Conservation of Nature and Natural Resources. It is threatened by habitat loss due to logging and forest clearance for agriculture. Its dependence on high-canopy forests make it particularly susceptible to tree felling, and it is now rare or absent in those parts ofits distribution where there has been extensive logging that breaks up the canopy. The Black Colobus is also widely hunted forits meat and skin, with populations on Bioko Island being under considerable pressure. There are no estimates ofthe total population of Black Colobus, but it is now present only in protected or undisturbed inaccessible areas. The largest numbers are thought to be in Lopé National Park, Gabon, where there is a population of 50,000 - 55,900 individuals. Further data on the distribution and status of the Black Colobus are required. More effective protection is also needed against hunting and habitat loss. The Bioko Black Colobus has undergone the biggest decline of all Bioko’s primates — more than 60 % from 1986 to 2006. It was common on Pico Basilé in 1986 but has been heavily hunted since 2000. It is becoming increasingly rare outside the Gran Caldera and Southern Highlands. It occurs in Basilé National Park and Gran Caldera and Southern Highlands Scientific Reserve (Bioko). The Gabon Black Colobus has fared somewhat better butis still declining, largely because of habitat loss and hunting. In mainland Equatorial Guinea, for example, its distribution was reduced to one-third ofits former size by 1967. It occurs in Dipikar and Douala-Edéa reserves in Cameroon, Monte Alen National Park in Equatorial Guinea, and the Lopé and the Gamba complex of protected areas in Gabon. The Gabon Black Colobus is also found in Camp Ma’an National Park and Dja Biosphere Reserve in Cameroon and Odzala National Park in the Republic of the Congo.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8BFF8EFAFE678CF72AF3D2.taxon	materials_examined	Afrique.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8BFF8EFAFE678CF72AF3D2.taxon	discussion	A form known as “ Dollman’s Colobus ” dollmani (a cross of C. vellerosus x C. polykomos, although phenotypically much closer to C. vellerosus) formerly occurred in an area between the Sassandra River and the Nzi-Bandama River system in Ivory Coast as a hybrid swarm. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8BFF8EFAFE678CF72AF3D2.taxon	distribution	Distribution. E Ivory Coast (E of Nuzi-Bandama River system), S Burkina-Faso, Ghana, Togo, Benin, and SW Nigeria.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8BFF8EFAFE678CF72AF3D2.taxon	description	Descriptive notes. Head-body 56 — 64 - 5 cm (males) and 58 - 66 cm (females), tail 81 - 97 cm; weight 8: 4.9 - 9 kg (males) and 7.8 - 3 kg (females). The White-thighed Colobus is largely black, with a broad, bushy, snowy-white circumfacial ruff (not extending back onto the crown or far back under the throat) and either no epaulettes or just a few white hairs on shoulders. There is no whorl on the crown. Tail is not as long as that of the King Colobus (C. polykomos), being somewhat less than 160 % of head — body length; it is also entirely white, with a slight tuft. There is a white patch on each upper thigh that extends all around ischial callosities in males and around ventral and lateral margins in females. Hair on the throat is thick. Nose is long and convex. In its cranium and mtDNA, the White-thighed Colobusis very similar to the King Colobus, but the loud call of the male is derived and shared with the Guereza (C. guereza).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8BFF8EFAFE678CF72AF3D2.taxon	biology_ecology	Habitat. L. owland rainforest and deciduous gallery, swamp, semi-deciduous, gallery, and dry forest. In the northern part of its distribution, the White-thighed Colobusis also found in forest patches in savanna areas, where groups have been known to travel across open ground between trees.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8BFF8EFAFE678CF72AF3D2.taxon	food_feeding	Food and Feeding. The White-thighed Colobus eats mainly young leaves, along with mature leaves, seeds, and fruit pulp.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8BFF8EFAFE678CF72AF3D2.taxon	breeding	Breeding. Female White-thighed Colobus lack a perineal sexual swelling, and infants are born with a white coat that darkens to the adult pattern within a few months.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8BFF8EFAFE678CF72AF3D2.taxon	activity	Activity patterns. White-thighed Colobus are diurnal and arboreal. At Boabeng-Fiema Monkey Sanctuary, J. Teichroeb and colleagues found them to spend 59 % of their time resting, 24 % feeding, 14 % moving, and 3 % engaging in social behavior.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8BFF8EFAFE678CF72AF3D2.taxon	biology_ecology	Movements, Home range and Social organization. Daily movements of the Whitethighed Colobus are 250 - 367 m, with home ranges of 1148 ha. Group sizes vary from two to 33 individuals, with multiple adult males and females. Dispersal is sex-biased,; males leave their natal group, but females are also known to disperse.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8BFF8EFAFE678CF72AF3D2.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Vulnerable on The IUCN Red List. The White-thighed Colobusislisted as Class A in the African Convention on the Conservation of Nature and Natural Resources. It is threatened primarily by hunting and secondarily by habitat loss. The region it inhabits has a dense and rapidly growing human population, forest destruction has been extensive, and wildlife hunting is uncontrolled in most places. There has been no systematic survey of the White-thighed Colobus, but it is known to have become rare in several national parks in Ghana where it had been assumed to be secure. It is now also very rare in Benin, Togo, and Nigeria. It occurs in 18 protected areas: Bia, Bui, Digya, Mole, and Nini-Suhien national parks, Boabeng-Fiema Monkey Sanctuary, Bomfobiri Wildlife Sanctuary, Ankasa and Kalakpa game reserves, Kogyae Strict Nature Reserve and Willi Falls Reserve in Ghana; Comoe National Park in Ivory Coast; Kainji Lake National Park, Igangan Ohosu and Olokemeji forest reserves and Upper Ogun game reserve in Nigeria; and Fazao-Malfakassa National Park and Keran Reserve in Togo. It is most secure in Boabeng-Fiema Monkey Sanctuary where monkeys are considered sacred by the local community and are not hunted.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8CFF89FF3E64A2F5B2FE1D.taxon	materials_examined	Sierra Leone (probably Sherbro Island).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8CFF89FF3E64A2F5B2FE1D.taxon	discussion	Modern taxonomic arrangements of the colobus monkeys either divide the red colobus and the Olive Colobus into two genera, Piliocolobus and Procolobus, respectively, or consider them to belong to one genus, Procolobus, with two subgenera (Procolobus for the Olive Colobus and Pilwcolobus for the red colobus). We follow here C. P. Groves in his publications of 2001 and 2007 in using two genera. The exact boundary between P. badius and P. temminckii is unclear, but the two are believed to be geographically separated. P. badius and P. waldronae meet at the Bandama River, Ivory Coast. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8CFF89FF3E64A2F5B2FE1D.taxon	distribution	Distribution. Sierra Leone (W limit Rokupr, 9 ° N, 12 ° 34 ’ 48 ” W), S Guinea, Liberia, and W Ivory Coast (E to the Nzi-Bandama River system).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8CFF89FF3E64A2F5B2FE1D.taxon	description	Descriptive notes. Head-body 50 - 63 cm (males) and 43 - 63 cm (females), tail 63 — 77 cm (males) and 63 - 77 cm (females); weight 9 - 12.5 kg (males) and 6 - 9 kg (females). These measurements, based mainly on animals from Ivory Coast, reveal some difference in size between males and females, but weights for Sierra Leone specimens show no such dimorphism: males 8 - 3 kg (6.4 - 9.6 kg) and females 8 - 2 kg (7 - 10 kg). Body of the Upper Guinea Red Colobus is a deep shining black, the tone extending to upper parts of limbs and forward to the forehead. Undersides, including inner surfaces of limbs, all of lower limb segments, and cheeks, are a deep reddish maroon. Pubic area and hind parts of thighs are white. Tail is mostly red, but it becomes darker toward the tip. Face is wrinkled, nearly black, but pink (depigmented) around eyes and mouth. Nostrils are raised on a prominentfleshy base. Skull is broad with a broad palate and short face. Adult female Upper Guinea Red Colobus have a small clitoris, and periodic sexual swellings are large. Infants are gray above and whitish below, with red tones on hands, feet, and tail.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8CFF89FF3E64A2F5B2FE1D.taxon	biology_ecology	Habitat. Primary or mature old growth moist forest. The Upper Guinea Red Colobusis more dependent on to tall forest than Temminck’s Red Colobus (P. temminckii).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8CFF89FF3E64A2F5B2FE1D.taxon	food_feeding	Food and Feeding. The Upper Guinea Red Colobus eats young leaves, seeds, mature leaf parts, fruits, and flowers.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8CFF89FF3E64A2F5B2FE1D.taxon	breeding	Breeding. Although somewhat variable, the female Upper Guinea Red Colobus does display a substantial pink sexual swelling during the periovulatory period. Breeding occurs throughout the year. The interbirth interval is ¢. 24 months. A single young is born after a gestation of 198 days. Neonates are the same general color as adults but slightly lighter (i. e. grayish on the back with a whitish ventral surface and reddish or orange tints on sides, tail, limbs, and extremities). Subadult males have a large red perineal organ around the anus between the ischial callosities.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8CFF89FF3E64A2F5B2FE1D.taxon	activity	Activity patterns. The Upper Guinea Red Colobus is diurnal and arboreal. At Tiwai Island Wildlife Sanctuary, groups spent c. 55 % their day feeding, 37 % resting, 5 % moving, and 3 % engaging in other activities.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8CFF89FF3E64A2F5B2FE1D.taxon	biology_ecology	Movements, Home range and Social organization. Daily movements of the Upper Guinea Red Colobus are ¢. 822 - 922 m, and groups occupy home ranges of 50 - 65 ha. Group size is 33 - 90 individuals, with multiple adult males and females. Males stay in their natal groups, and females typically disperse. Females spend more time with males than they do with other females. They groom males and other females equally and support males in conflict. According to A. Korstjens’s study, males groom infrequently and do not support females in conflicts. Upper Guinea Red Colobus suffer from predation by Chimpanzees (Pan troglodytes), and they travel with Diana Monkeys (Cercopithecus diana) as a way of mutually enhancing their ability to detect Chimpanzees and other predators such as crowned hawk-eagles (Stephanoaetus coronatus).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8CFF89FF3E64A2F5B2FE1D.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Endangered on The IUCN Red List (as Procolobus badius). The Upper Guinea Red Colobusis listed as Class B in the African Convention on the Conservation of Nature and Natural Resources. It is threatened mainly by habitat loss and hunting. Deforestation through logging, charcoal production, and clearance for agricultural land, including plantations, has occurred over much of its distribution, especially in the last century. Subsistence and commercial hunting have severely affected populations of the Upper Guinea Red Colobus. Improved access to remote forest via logging roads has increased hunting. Much ofits distribution has been impacted by civil conflict since 1989, and it is not yet clear to what extent this has affected populations in Sierra Leone, Liberia, and Ivory Coast. The Upper Guinea Red Colobus occurs in Tai Forest National Park in Ivory Coast; Sapo National Park in Liberia; and Outamba-Kilimi National Park, Gola Forest Reserve, and Tiwai Island Wildlife Sanctuary in Sierra Leone. Hunting has reportedly extirpated it in four forest reserves and Marahoué National Park between the Sassandra and Bandama rivers in Ivory Coast.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8CFF8AFA38673EFD14F81B.taxon	materials_examined	type locality unknown.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8CFF8AFA38673EFD14F81B.taxon	discussion	Modern taxonomic arrangements of the colobus monkeys either divide the red colobus and the Olive Colobus into two genera, Piliocolobus and Procolobus, respectively, or consider them to belong to one genus, Procolobus, with two subgenera (Procolobus for the Olive Colobus and Piliocolobus for the red colobus). We follow here C. P. Groves in his publications of 2001 and 2007 in using two genera. The exact boundary between P. badius and P. temminckii is unclear, but the two are believed to be geographically separated. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8CFF8AFA38673EFD14F81B.taxon	distribution	Distribution. From the Gambia and Upper Senegal rivers through SW Senegal, Gambia, and Guinea-Bissau (absent E of Rio Corubal, except for a supposed sighting at Catio, near the border with Guinea) to NW Guinea (E to Kopulau Camp, 12 ° 24 ° N, 13 ° 34 ° 48 ” W); possibly in NW Sierra Leone. Although reported from the Fouta Djallon Highlands in Guinea, there are no published records available to confirm this.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8CFF8AFA38673EFD14F81B.taxon	description	Descriptive notes. Body measurements are not available but museum records suggest Temminck’s Red Colobus is similar in size to (perhapsa little smaller than) the Upper Guinea Red Colobus (P. badius). It is medium gray dorsally (extending to upper parts of limbs), with flanks and limbs, and a collar round the neck, light orange-red, becoming whitish on the undersides. Thereis a triangular patch of white fur on the buttocks. Fingers and toes darken to rufous. Face is dark with rings of pink encircling the eyes, and nose is broad and upturned, raised on a fleshy base as in the Upper Guinea Red Colobus. Scrotum of Temminck’s Red Colobus appears to be less pendulous than in the Upper Guinea Red Colobus, and the perineal organ is very small. It appears that the typical Temminck’s Red Colobusis confined to areas of Gambia and Senegal north of the Gambia River, and south of this river, there is a gradual darkening and loss of the red collar and white strip on the underside. There seems to be a fairly gradual transition between this pale colored dry-forest form and the typical deeply colored rainforest form. In a mtDNA tree, N. Ting found Temminck’s Red Colobus nested among the Upper Guinea Red Colobus, with whom it also shares very similar vocalizations.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8CFF8AFA38673EFD14F81B.taxon	biology_ecology	Habitat. Dry deciduous and gallery forest, usually near open bodies of water, in Senegal. Temminck’s Red Colobus is unique among red colobus in that it often lives in savanna woodland. In the 1970 s, A. Galat-Luong and G. Galat reported that the northernmost population in Saloum (Senegal) lived only in tropical, dense high forest, but with widespread degradation and loss of these forests, in recent years, Temminck’s Red Colobus have begun to occupy more open habitats, such as degraded remnants of gallery forests with interrupted canopies, and even true wooded savanna, mangrove swamps, and farmland. Researchers in Saloum in the 1970 s never found Temminck’s Red Colobus away from tall dense forest, but with forest loss and degradation by the 1990 s, they were found in open woodland habitat and even mangroves more than one-third of the time, often associating with the open woodland Green Monkeys (Chlorocebus sabaeus). As forest dwellers, Temminck’s Red Colobus would generally travel and forage ¢. 10 m above the ground. In more damaged forests and open habitats, their average height above the ground can drop by nearly half (a little more than 5 m), and they actually go to the ground more often. In the Fathala Reserve of the Saloum Delta National Park in coastal Senegal, changes in diet, tendencies toward terrestriality and associations with other primates (particularly Green Monkeys), use of open habitats, and use of mangrove swamps seem to be successive accommodations to loss of their preferred forests.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8CFF8AFA38673EFD14F81B.taxon	food_feeding	Food and Feeding. Temminck’s Red Colobus eat young leaves and leaf buds, along with fruits and seeds. Mature leaves, flowers, and flower buds are also consumed. Diets of six groups, studied by B. Gatinot in the Fathala Reserve in 1973 - 1974, contained 21 - 8 % young leaves, 20 - 1 % mature leaves, 16 - 5 % buds, 14 - 4 % flowers, 9 - 5 % fruits, and 11 - 4 % other items. Diets included 24 species of plants. D. Diouck also carried out a dietary study of two groups in Fathala in 1994 - 1996. Diets had changed considerably from the 1973 - 1974 study and included more young leaves and fruits, fewer mature leaves and buds, and considerably fewer flowers: 56 - 7 % young leaves, 18 -: 8 % fruits, 13 - 4 % buds, 6 - 3 % mature leaves, 0 - 8 % flowers, and 4 % other items. The number of plant species in the diet had dropped from 24 to 17. This change was associated with changes in Fathala during this period; c. 75 % of the gallery forest where they had been studied in the 1970 s had been lost, and the forest that remained was largely degraded, with broken canopies and a 30 % reduction in the diversity of woody plants. Temminck’s Red Colobus spent more time on or near the ground and included more herbs, grasses, grass seeds, and groundnuts in their diets.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8CFF8AFA38673EFD14F81B.taxon	breeding	Breeding. Female Temminck’s Red Colobus display large sexual swellings during the periovulatory period. A single young is born after a gestation of 173 - 174 days. Neonates are the same general color as adults but slightly lighter (light beige or gray with a darker back and a bit of orange on the tail).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8CFF8AFA38673EFD14F81B.taxon	activity	Activity patterns. Temminck’s Red Colobusis diurnal and arboreal, but it can become increasingly terrestrial when adapting to reduced and degraded forest habitats (e. g. being forced to travel and forage in savanna and woodland environments surrounding gallery forest). At Abuko, it rested for 52 % of the day, fed for 21 %, traveled for 13 %, and engaged in social activities for 13 %; the remaining 1 % was dedicated to a variety of other activities.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8CFF8AFA38673EFD14F81B.taxon	biology_ecology	Movements, Home range and Social organization. Home range sizes of Temminck’s Red Colobus are 11 - 34 ha. Group sizes typically are 12 - 28 individuals, but there has been evidence of exceptionally large groups (e. g. up to 62 individuals). Groups typically have multiple adult males and females, and both sexes disperse. D. Starin found that females groom mostly among themselves, and males only infrequently groom others. Curiously, Temminck’s Red Colobus may also groom Green Monkeys when they temporarilyjoin their groups after dispersing from their natal groups. Predators recorded include African rock python (Python sebae) and, when near to or on the ground, dogs and Spotted Hyenas (Crocuta crocuta).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8CFF8AFA38673EFD14F81B.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Endangered on The IUCN Red List (as Procolobus badius temminckii). Temminck’s Red Colobusis listed as Class B in the African Convention on the Conservation of Nature and Natural Resources. Its major threat is habitat loss due to forest conversion by agriculture, overgrazing, fires, and tree-cutting, combined with decreasing rainfall in the part of West Africa where it occurs. Although hunting is also a threat to Temminck’s Red Colobus (particularly in the southern part ofits distribution), it appears to be somewhat less affected than Miss Waldron’s Red Colobus (P. waldronae) and the Upper Guinea Red Colobus. In The Gambia, monkeys are hunted more as farm pests than for consumption as meat, and red colobus are minor farm pests compared with baboons, Green Monkeys, and Patas Monkeys (Erythrocebus patas). In Senegal, there are probably fewer than 400 - 500 Temminck’s Red Colobus surviving in Saloum Delta National Park and probably fewer than 100 in the isolated Niokolo-Koba National Park and north-western Guinea. They also occur in Abuko and River Gambia national parks in Gambia, Cufada National Park in Guinea-Bissau, and Badiar National Park in Guinea. They may have been extirpated from Basse Casamance National Park in Senegal. The absence of many large and well-managed protected areas means that the overall population of Temminck’s Red Colobusis likely to continue to decline, particularly in the southern part ofits distribution where civil conflict has persisted and hunting pressure is increasing.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8FFF8AFF996D23F97CFABA.taxon	materials_examined	Goaso, Ashanti, Ghana.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8FFF8AFF996D23F97CFABA.taxon	discussion	The name waldroni is here corrected to waldronae, as required by Article 31.1.2 of the International Code of Zoological Nomenclature, 4 " edition, published in 1999. Modern taxonomic arrangements of the colobus monkeys either divide the red colobus and the Olive Colobus into two genera, Piliocolobus and Procolobus, respectively, or consider them to belong to one genus, Procolobus, with two subgenera (Procolobus for the Olive Colobus and Piliocolobus for the red colobus). We follow here C. P. Groves in his publications of 2001 and 2007 in using two genera. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8FFF8AFF996D23F97CFABA.taxon	distribution	Distribution. SE Ivory Coast; possibly in SW Ghana.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8FFF8AFF996D23F97CFABA.taxon	description	Descriptive notes. Head-body 45 - 67 cm (males) and 42 - 56 cm (females), tail 56 68 cm (males) and 52 - 67 cm (females); weight 5 - 5 — 6 - 3 kg. Miss Waldron’s Red Colobus is, on average, considerably smaller than the Upper Guinea Red Colobus (FP. badius). There is a low degree of sexual dimorphism, and its tail is shorter. It is characterized by its very black limbs and flat nose (without a raised fleshy base). Crown, nape, shoulders, and back are the same shining black as in the Upper Guinea Red Colobus, but forehead, limbs, whiskers, lower half of flanks, and underside are bright maroon, and hind parts of thighs and scrotum are white. Tail is black. Face is slaty-blue, with pink lips but no depigmentation around the eyes. According to mtDNA analysis on a single individual, Miss Waldron’s Red Colobusis not as closely related to the Upper Guinea Red Colobus as some ofits external features might suggest. It carries a mitochondrial lineage quite divergent from those sampled thus far in the Upper Guinea Red Colobus and Temminck’s Red Colobus (P. temminckiz).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8FFF8AFF996D23F97CFABA.taxon	biology_ecology	Habitat. Moist forest zone, although parts contained forest that had a marked dry season and semi-deciduous trees. Miss Waldron’s Red Colobus uses the upper levels of the forest canopy, above the sympatric King Colobus (Colobus polykomos).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8FFF8AFF996D23F97CFABA.taxon	food_feeding	Food and Feeding. The diet of Miss Waldron’s Red Colobus is similar to that of the Upper Guinea Red Colobus and contains mainly leaves, supplemented by fruits, seeds, and flowers.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8FFF8AFF996D23F97CFABA.taxon	breeding	Breeding. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8FFF8AFF996D23F97CFABA.taxon	activity	Activity patterns. Miss Waldron’s Red Colobus is diurnal and arboreal.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8FFF8AFF996D23F97CFABA.taxon	biology_ecology	Movements, Home range and Social organization. A short study of Miss Waldron’s Red Colobus at Bia National Park in Ghana in the 1970 s recorded a group of 34 individuals in a home range of 140 ha and another group of 58 individuals in a home range of 100 ha. Its vocalizations are similar to those of the Upper Guinea Red Colobus.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8FFF8AFF996D23F97CFABA.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Critically Endangered on The IUCN Red List (as Procolobus badius waldronae). Miss Waldron’s Red Colobusis listed as Class B in the African Convention on the Conservation of Nature and Natural Resources. In the past, it ranged from the Nzi-Bandama River system in south-eastern Ivory Coast to south-western Ghana (Bibianha, 6 ° 17 ° N, 2 ° 4 ’ 48 ” W), not quite reaching the Volta River. It has been extirpated over most of its distribution. There have been no confirmed sightings since 1978, and surveys carried out from 1993 to the present have yet to confirm the survival of any living individuals. Accordingly, in 2000, Miss Waldron’s Red Colobus was declared probably extinct. Since then, however, there have been a few unconfirmed sightings (in addition to a photograph) reported from the Ehy Forest, Ivory Coast, where a skin and tail were discovered in the possession of a hunter in 2002. Calls of what are believed to be Miss Waldron’s Red Colobus in this region were heard in early 2008, giving some hope that the taxon may yet persist in the wild. Further surveys are urgently needed. Massive habitat loss and hunting pressure are responsible for its decline.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8FFF8BFAE06082FAAFF892.taxon	materials_examined	Tanzania, Zanzibar.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8FFF8BFAE06082FAAFF892.taxon	discussion	Modern taxonomic arrangements of the colobus monkeys either divide the red colobus and the Olive Colobus into two genera, Piliocolobus and Procolobus, respectively, or consider them to belong to one genus, Procolobus, with two subgenera (Procolobus for the Olive Colobus and Piliocolobus for the red colobus). We follow here C. P. Groves in his publications of 2001 and 2007 in using two genera. P. kirkii is the easternmost representative of the genus. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8FFF8BFAE06082FAAFF892.taxon	distribution	Distribution. Zanzibar I (mainly in the SE) and Muyuni and Uzi Is, off E Tanzania. Introduced on nearby Pemba I.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8FFF8BFAE06082FAAFF892.taxon	description	Descriptive notes. Head-body 45 - 50 cm, tail 42 - 55 cm; weight 7 - 9 kg. It is possible that female Zanzibar Red Colobus are larger than males. It is the smallest and most distinctive species of Piliocolobus. Hindlimbs are elongated. It is an essentially tricolored form; arms, shoulders, and forepart of the back are black, with the remainder of the back, crown, and base of the tail brick-red. Shaggy hair surrounds the face like a halo. Underside (including inner surfaces of forelimbs and all of hindlimbs) and terminal part of the tail are creamy-white. Hands and feet are black and fringed with white hairs. Face is black with a pink zone around the mouth and lower part of the nose; this pattern occurs in juveniles of many other red colobus, but it is retained for life only in Zanzibar Red Colobus. Eyes are ruby-red. Skull has a short facial skeleton but a long palate. A mtDNA places the Zanzibar Red Colobus in a distinct East African clade with the Udzungwa Red Colobus (P. gordonorum), with which it shares very similar vocalizations.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8FFF8BFAE06082FAAFF892.taxon	biology_ecology	Habitat. Secondary forest patches, swamp forest, mangrove, fallow bush, and agricultural land, from sea level to 110 m. The Zanzibar Red Colobus is found primarily in areas of groundwater forest and scrub forest on coral rag on the southern and eastern side of the island. It originally inhabited tropical evergreen forest (now largely destroyed). It prefers the forest canopy.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8FFF8BFAE06082FAAFF892.taxon	food_feeding	Food and Feeding. Young leaves and leaf buds account for more than 50 % of the diet of the Zanzibar Red Colobus. In forest-dwelling groups, unripe fruit accounts for an additional 25 % of the diet, whereas in groups living in agricultural areas, leaves from herbaceous species are the second-most consumed item (c. 15 % of the annual diet). Some groups frequently feed on mangrove leaves, and groups living on coral rag appear to subsist on a drier, coarser diet than any recorded for red colobus monkeys. Individual Zanzibar Red Colobus occasionally eat soil or insect clay, and also charcoal from a burned stump.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8FFF8BFAE06082FAAFF892.taxon	breeding	Breeding. Female Zanzibar Red Colobus display a substantial pink sexual swelling during the periovulatory period. Births take place throughout the year, and a single young is born after a gestation of ¢. 5 months. The interbirth interval is 26 - 55 months. The neonatal coat is white (not blackish as in other red colobus).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8FFF8BFAE06082FAAFF892.taxon	activity	Activity patterns. The Zanzibar Red Colobus is diurnal and arboreal. Studying groups at two differentsites, K. Siex found they spent 44 - 47 % of their time resting, 29 % feeding, 6 - 12 % traveling, 7 - 15 % engaging social activities, and 7 - 15 % in other activities.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8FFF8BFAE06082FAAFF892.taxon	biology_ecology	Movements, Home range and Social organization. Daily movements average 310 - 565 m, and home ranges are 13 - 25 ha (home ranges overlap). The Zanzibar Red Colobus lives in multimale-multifemale groups, averaging 30 individuals.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8FFF8BFAE06082FAAFF892.taxon	conservation	Status and Conservation. CITES Appendix I. Classified as Endangered on The IUCN Red List (as Procolobus kirkii). The Zanzibar Red Colobusis listed as Class A in the African Convention on the Conservation of Nature and Natural Resources. It is nominally protected in Tanzania under the Forest Resource Management and Conservation Act of 1996. These beautiful monkeys are threatened by hunting and land clearance. Two specimens of the Zanzibar Red Colobus obtained in the mid- 19 " century reportedly came from the Pangani River, on mainland Tanzania, but it is certainly extirpated there now. Vast tracts of Zanzibar’s native forest have been destroyed for timber, charcoal production, cultivation, and bush-burning. Local people tolerate the Zanzibar Red Colobus and seem to appreciate its importance as a tourist attraction; however, it is occasionally shot for food or sport, or as a crop pest. On Zanzibar, it is found mainly in the south-eastern part of the island in Jozani-Chwaka Bay National Park, adjacent agricultural areas to the south, and the coral thickets and mangrove swamps of Uzi Island, 10 km to the south-west. It is also found at low densities in isolated populations in coral thickets along Zanzibar’s eastern coast from Kiwengwa in the north to Mnyambiji in the south. On the western coast, there is a small isolated group in mangrove swamps of Maji Mekundu. While the Zanzibar Red Colobus has adapted well to human encroachment, its extent of occurrence is less than 5000 km * and probably less than 760 km?, and there is severe fragmentation and continuing declines in area of occupancy, habitat, and numbers of mature individuals. Less than 50 % of the population is legally protected and reasonably secure in the small Jozani Chwaka Bay National Park (c. 60 km ®). There are occasional deaths due to road kills south of the park, but the installation of speed bumps at Jozani has reduced these incidences. The highest densities occur in ¢. 40 ha of agricultural land adjacent to the southern border ofJozani Chwaka Bay National Park (784 ind / km? in 1999), but they are due to population compression following the destruction of adjacent habitat rather than intrinsic growth of the population. The density in coral thickets adjacent to Jozani Forest is probably less than 50 ind / km? The Jozani subpopulation was thought to contain ¢. 500 individuals, and in 2008, the total population was estimated at less than 2000 individuals. A small translocated population (c. 56 individuals) also occurs in the Masingini Forest Reserve. Creation of a new protected area at Kiwengwa, for the protection of remaining patches of coral thicket in the south and corridors between remaining habitat patches, is urgently needed for survival of the Zanzibar Red Colobus. About 14 individuals were introduced to Ngezi Forest Reserve, Pemba Island, around 1974, and a small population persists there today.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8EFF8BFFD862A6F745F6B4.taxon	materials_examined	Tanzania, Uzungwa (= Udzungwa) Mountains, Uhehe.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8EFF8BFFD862A6F745F6B4.taxon	discussion	Modern taxonomic arrangements of the colobus monkeys either divide the red colobus and the Olive Colobus into two genera, Piliocolobus and Procolobus, respectively, or consider them to belong to one genus, Procolobus, with two subgenera (Procolobus for the Olive Colobus and Piliocolobus for the red colobus). We follow here C. P. Groves in his publications of 2001 and 2007 in using two genera. There is a rare red-backed morph (representing c ¢. 2 % of the population), in which the upper bodyis black with a brick-red lumbar region and rump. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8EFF8BFFD862A6F745F6B4.taxon	distribution	Distribution. SW Tanzania, known only from the Udzungwa Mts, a few small tracts of forest between the Little Ruaha and Ulanga rivers (S of Iringa), and several forest patches of the Kilombero Valley at the E base of the Udzungwa Mts (Magombera, Kiwanga, Kalunga, and in gallery forest along the Msolwa River S of the railway to its confluence with the Kilombero River, including the Ibiki and Msitu ya Kinjugu forests).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8EFF8BFFD862A6F745F6B4.taxon	description	Descriptive notes. Head-body 60 - 66 cm (females), tail 64 - 69 cm (females); no measurements for males or body weight data for either sex are available. The Udzungwa Red Colobus is somewhat larger, but with a shorter tail, than the Ashy Red Colobus (P. tephrosceles). It is a deep shining black above (rarely with a red lumbar region and rump), with white on underside and inner surfaces of limbs. Arms and thighs are black, but shanks are mixed with silvery. Lower legs black mixed with silvery. Tail is short and slightly bushy, being black or mixed with ocherous above and tending to be off-white below. Crown is a shiny red, with laterally directed hairs forming a sort of toupee. Face lacks pigmentation around the nose and mouth, and cheeks are white and separated from red of the crown by a thick black line along the temples. A “ panache ” (a tuft of hair) is present but weakly developed. Skull is broad, short-faced, and small-toothed. A mtDNA analysis shows that the Udzungwa Red Colobus is most closely related to the Zanzibar Red Colobus (P. kirkii), but distinct.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8EFF8BFFD862A6F745F6B4.taxon	biology_ecology	Habitat. A variety of forest habitats, including lowland old growth, riparian, groundwater, secondary, and montane forests of the Udzungwa Mountains and Miombo (Brachystegia, Fabaceae) woodland adjacent to old growth or secondary forest in the lower parts of the Mwanihana Forest in the Udzungwa Mountains National Park and elsewhere. Systematic population surveys examining the relative abundance of primates in the Mwanihana Forest found that the Udzungwa Red Colobus preferred moderate to old growth semi-deciduous forest, but it was also found nearly as often in evergreen forest. It was recorded in deciduous forest and open areas (logged, regenerating forest with shrubs and scattered trees) but in lower numbers. The Udzungwa Red Colobus occurs at elevations of 200 - 2200 m, althoughit is less abundant at the higher elevations.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8EFF8BFFD862A6F745F6B4.taxon	food_feeding	Food and Feeding. Diet of the Udzungwa Red Colobus includes young leaves and mature leaves, fruits, flowers, buds, and possibly seeds.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8EFF8BFFD862A6F745F6B4.taxon	breeding	Breeding. Female Udzungwa Red Colobus display a substantial pink sexual swelling during the periovulatory period (larger than in the Ashy Red Colobus).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8EFF8BFFD862A6F745F6B4.taxon	activity	Activity patterns. There is no specific information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8EFF8BFFD862A6F745F6B4.taxon	biology_ecology	Movements, Home range and Social organization. The movements and home range of the Udzungwa Red Colobus are unknown, but group size is highly variable at 3 - 83 individuals. Groups are smaller in forests that are small (less than 5 km?), dominated by deciduous trees, or heavily degraded by human activities than are groups living in large, intact, and relatively mature blocks of moist, mixed evergreen and semi-deciduous forest. Groups typically have multiple adult males and females, and variation in numbers of adult males accounts for much of the variability in group size.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8EFF8BFFD862A6F745F6B4.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Endangered on The IUCN Red Last (as Procolobus gordonorum). The Udzungwa Red Colobusis listed as Class B in the African Convention on the Conservation of Nature and Natural Resources. It has an extent of occurrence of less than 5000 km ®, populations are highly fragmented with no or limited movement between forest fragments, and there is a continuing decline in the distributional extent of occurrence and area of occupancy due to logging, conversion to agriculture, collection of firewood, and charcoal production. The Udzungwa Red Colobus is sometimes hunted for its meat, particularly in the western and south-western parts of the range. Total population size is unknown and difficult to determine, but clearly in the thousands. At least one published estimate puts it at 15,400 individuals, but this might be too high. An earlier estimate by W. Rodgers and K. Homewood in 1982 was 10,000 individuals, but they emphasized that the largest single interbreeding population was certainly less than 2000. In the Udzungwa Mountains, the Udzungwa Red Colobus was the most frequently seen primate, with an average of 0 - 59 groups / km walked. It occurs in ten protected areas in Tanzania: Udzungwa Mountains National Park; Kiranzi-Kitungulu, Magombero, Mangula, Matundu, New Dabaga / Ulangambi, Nyanganje, Udzungwa Scarp, and West Kilombero forest reserves; and Selous Game Reserve. It is relatively secure in Udzungwa Mountains National Park, but only about half ofits range is protected in this area. Its remaining habitat is either on public land or in forest reserves (including New Dabaga / Ulangambi Forest Reserve) and is not effectively protected. Magombero Forest Reserve, for example, still needs to be legally incorporated into Selous Game Reserve, which the government of Tanzania had promised to do in 1980. In 2004, T. Struhsaker and colleagues highlighted the following conservation priorities for the Udzungwa Red Colobus: 1) annexation of a number of forest reserves to the Udzungwa Mountains National Park (West Kilombero, Udzungwa Scarp, Matundu, and Nyanganje); 2) annexation of a number of forests to Selous Game Reserve (Magombera, Ibiki, and all other forest remnants along the Msolwa River); 3) prevention of fire to allow forest regeneration and expansion and creation of corridors between forest blocks; and 4) more effective law enforcement to prevent fires, illegal logging, and poaching.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8EFF94FA256C80FB12FA40.taxon	materials_examined	Equatorial Guinea, Fernando Poo (Bioko) Island.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8EFF94FA256C80FB12FA40.taxon	discussion	Modern taxonomic arrangements of the colobus monkeys either divide the red colobus and the Olive Colobus into two genera, Piliocolobus and Procolobus, respectively, or consider them to belong to one genus, Procolobus, with two subgenera (Procolobus for the Olive Colobus and Piliocolobus for the red colobus). We follow here C. P. Groves in his publications of 2001 and 2007 in using two genera. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8EFF94FA256C80FB12FA40.taxon	distribution	Distribution. Bioko I (Equatorial Guinea), now mainly restricted to the SW.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8EFF94FA256C80FB12FA40.taxon	description	Descriptive notes. Head-body c. 62 cm (males) and c. 64 cm (females), tail ¢. 60 cm (males) and c. 66 cm (females); weight 9 kg (males) and c. 5 - 8 kg (females). Pennant’s Red Colobusis a rather large, short-tailed species. Photographs taken in the wild suggest that adult males are large and robustly built. In many respects, Pennant’s Red Colobusis similar to Bouvier’s Red Colobus (P. bouvieri), but it is darker, with speckled red and black, almost a solid black, on shoulders, becoming bright red on flanks. Underparts are pale reddish-white to white. Hands and feet are contrastingly black, this tone extending some way up on limbs. Tail is black above and deep red below. Cheek whiskers are white, and crown is black. There is a prominent tuft at the base of each ear. Face is black. Scrotum is not conspicuous, and the clitoris is prominent. Teeth are relatively small, skull is generally narrow, facial skeleton is long, and muzzle is broad. A mtDNA study placesit close to its mainland neighbor, Preuss’s Red Colobus (P. preussi), but its vocalizations are distinct.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8EFF94FA256C80FB12FA40.taxon	biology_ecology	Habitat. [. ow and montane forest from sea level to at least 1500 m, and perhaps even 1800 m. Southern forests on Bioko Island, where Pennant’s Red Colobus is most common, receive very abundant rainfall, some areas exceeding 10,000 mm / year.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8EFF94FA256C80FB12FA40.taxon	food_feeding	Food and Feeding. Diets include young leaves and mature leaves, fruit, flowers, buds, and possibly seeds.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8EFF94FA256C80FB12FA40.taxon	breeding	Breeding. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8EFF94FA256C80FB12FA40.taxon	activity	Activity patterns. Pennant’s Red Colobusis diurnal and arboreal.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8EFF94FA256C80FB12FA40.taxon	biology_ecology	Movements, Home range and Social organization. Pennant’s Red Colobus has never been studied in the wild. Its ranging behavior is unknown. Groups contain 5 - 30 individuals, with 1 - 3 adult males. Females travel alone and are probably the sex that disperses from natal groups. Humans are the only predators on Bioko Island; potential predators of red colobus on the mainland, such as crowned eagles (Stephanoaetus coronatus), Leopards (Panthera pardus), and Chimpanzees (Pan troglodytes), do not occur on Bioko Island.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF8EFF94FA256C80FB12FA40.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Endangered on The IUCN Red List (as Procolobus pennantii). Pennant’s Red Colobusis listed as Class B in the African Convention on the Conservation of Nature and Natural Resources. Its total distribution is less than 500 km? ®, and there are probably less than 5000 individuals remaining. In 2006, G. Hearn and colleagues estimated a 45 % decline in numbers between 1986 - 2006 due mainly to heavy hunting. It was estimated that more than 550 individuals were killed in 2004 alone. Pennant’s Red Colobus occurs in Gran Caldera and Southern Highlands Scientific Reserve. It maystill occur in Basilé National Park, but it has not recorded there since 1980 s. Shotgun hunting has been banned on Bioko Island, but the ban has not been strongly enforced.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF91FF94FFC66316F8E4FDCD.taxon	materials_examined	“ Gambia ” (selected by |. A. Allen in 1939). But according to R. E. Moreau and colleagues in 1946, “ Mongo, right bank of the Congo, two daysjourney by boat above the mouth of the Alima River. ”	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF91FF94FFC66316F8E4FDCD.taxon	discussion	Modern taxonomic arrangements of the colobus monkeys either divide the red colobus and the Olive Colobus into two genera, Piliocolobus and Procolobus, respectively, or consider them to belong to one genus, Procolobus, with two subgenera (Procolobus for the Olive Colobus and Piliocolobus for the red colobus). We follow here C. P. Groves in his publications of 2001 and 2007 in using two genera. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF91FF94FFC66316F8E4FDCD.taxon	distribution	Distribution. E Republic of the Congo in a small area on the right bank of the Congo River, from the mouth of the Likouala-aux-Herbes River, E of the Sangha River, to the lower reaches of Alima River, both tributaries of the lower Congo (c. 0 ° - 1 ° S). Records from Inoni, Lefini Faunal Reserve (3 ° S, 15 ° 30 ’ E), some distance S of the known distribution, require confirmation.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF91FF94FFC66316F8E4FDCD.taxon	description	Descriptive notes. Head — body ¢. 58 cm (females), tail ¢. 76 cm (females); no measurements for males or body weight data for either sex are available. Judging by measurements ofjust two specimens, Bouvier’s Red Colobus is a fairly small species, with a rather long-tail. Fur is glossy-red above, with a narrow, blackish, unbanded mantle extending to the middle of the back. Underparts are paler (partly brown and rufous), and thighs are yellower rufous like the flanks. Tail is dark at the root, turning goldenbrown to brownish-rufous distally. Crown is red and without whorls. There is a stuff black, superciliary band, a small tuft behind each ear, and a black brow-band bordering the forehead whorl, extending laterally onto the temples. Face is light slaty, with large pink eye-rings. The light face of Bouvier’s Red Colobus is quite unique; it varies from flesh-colored with blackish cheekbones and brows to darker with only nose and lips light colored. Those who have seen this form in the wild agree on the distinctiveness of the light face, which Rochebrune, in the type description, described as “ rosyflesh ” with blue-black cheekbones and eye-rings, and pale, rosy-tinted ears and black brows. Chin is covered with white hairs. Ischial callosities are pink. There are no DNA analyses for Bouvier’s Red Colobus.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF91FF94FFC66316F8E4FDCD.taxon	biology_ecology	Habitat. Swamp forest.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF91FF94FFC66316F8E4FDCD.taxon	food_feeding	Food and Feeding. Diets include young leaves and mature leaves, fruit, flowers, buds, and possibly seeds.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF91FF94FFC66316F8E4FDCD.taxon	breeding	Breeding. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF91FF94FFC66316F8E4FDCD.taxon	activity	Activity patterns. Bouvier’s Red Colobusis diurnal and arboreal.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF91FF94FFC66316F8E4FDCD.taxon	biology_ecology	Movements, Home range and Social organization. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF91FF94FFC66316F8E4FDCD.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Critically Endangered on The IUCN Red List (as Procolobus pennantii bouvieri). Bouvier’s Red Colobus is listed as Class B in the African Convention on the Conservation of Nature and Natural Resources. There appear to have been no reliable sightings of Bouvier’s Red Colobus since the 1970 s, and it may in fact be extinct. Heavy hunting for bushmeat and loss of habitat have caused declines of many wildlife species. Bouvier’s Red Colobus was at least formerly known to occur in the Lefini Faunal Reserve in the Republic of the Congo. Urgent investigations are needed to clarify the status of the very poorly known Bouvier’s Red Colobus.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF91FF94FAC16769F665F41F.taxon	materials_examined	Nigeria, Sampou-Apoi, 4 ° 55 ’ N, 6 ° 00 " E, 12 km SW of the junction of Sagbama and Egbedi Creeks, Niger Delta, Southern jaw Local Government camp area, Bayelsa State.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF91FF94FAC16769F665F41F.taxon	discussion	Modern taxonomic arrangements of the colobus monkeys either divide the red colobus and the Olive Colobus into two genera, Piliocolobus and Procolobus, respectively, or consider them to belong to one genus, Procolobus, with two subgenera (Procolobus for the Olive Colobus and Piliocolobus for the red colobus). We follow here C. P. Groves in his publications of 2001 and 2007 in using two genera. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF91FF94FAC16769F665F41F.taxon	distribution	Distribution. S Nigeria, apparently confined to the W part of the Niger Delta, between the Forcados-Nikrogha Creek and the Sagbama-Osiama-Agboi Creek (approximately the area S of 5 ° 09 ° N and W of 6 ° 07 ’ E); it may also occur sporadically farther E (almost to 7 ° E).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF91FF94FAC16769F665F41F.taxon	description	Descriptive notes. There are no measurements available. The Niger Delta Red Colobus is similar to Pennant’s Red Colobus (PF. pennantii) in having a black crown and a banded, black, and orange mantle, but whitish tone on underparts extends onto outer sides of arms, restricting orange-brown to a narrow band on trailing edges of outer arms or even completely excluding these tones. Black of hands and feet does not extend up the limbs. Tail is dark red-brown above, with a median dark brown dorsal band, chestnut or maroon below, terminal one-third to two-thirds of the tail becoming black-brown. There are patches of white on shoulders and cheeks, and black hairs over ears form conspicuous whorls. In mtDNA, a specimen of the Niger Delta Red Colobus seems to belong with East and Central African species, rather than with other West African species; its vocalizations follow a similar pattern.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF91FF94FAC16769F665F41F.taxon	biology_ecology	Habitat. Marsh forest.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF91FF94FAC16769F665F41F.taxon	food_feeding	Food and Feeding. Young leaves, leaf buds, and terminal parts of stems make up the majority of the diet of the Niger Delta Red Colobus. Other important food items include seeds, flowers and flower buds, and leaf petioles.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF91FF94FAC16769F665F41F.taxon	breeding	Breeding. Adult female Niger Delta Red Colobus develop swellings during the periovulatory period that are 12 cm long and 10 cm wide. Mating occurs throughout the year but most frequently during the rainy season in April-July. Small infants are seen most frequently during the dry season in December-March.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF91FF94FAC16769F665F41F.taxon	activity	Activity patterns. Niger Delta Red Colobus are diurnal and arboreal. In a study byJ. L. R. Werre, they spent 36 % of their time feeding, 33 % resting, 25 % traveling, and 6 % engaging in social behavior.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF91FF94FAC16769F665F41F.taxon	biology_ecology	Movements, Home range and Social organization. Daily movements of the Niger Delta Red Colobus are 450 - 1900 m, and home range is 73 ha. Groups contain 15 - 80 individuals, with multiple adult males and females. Females groom most frequently, and male grooming is typically directed toward females with perineal swellings.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF91FF94FAC16769F665F41F.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Critically Endangered on The IUCN Red List (as Procolobus pennantii epieni). The Niger Delta Red Colobusis listed as Class B in the African Convention on the Conservation of Nature and Natural Resources. It was only discovered in 1993 and was quite common within its limited distribution in the mid- 1990 s. More recent reports suggest rapid declines as a result of heavy hunting, compounded by habitat degradation. Artisanal-scale logging has seriously degraded the habitat, removing in the process many important food trees such as Hallea ledermannii (Rubiacae). The Niger Delta Red Colobus is not found in any protected areas, although it does occur in the proposed Apoi Creek Forest Reserve. Protection of its remaining habitat is an urgent priority, although civil conflict has been a hindrance to the implementation of effective conservation plans.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF91FF95FAC5693BFDB7F755.taxon	materials_examined	Cameroon, Barombi, Elephant Lake.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF91FF95FAC5693BFDB7F755.taxon	discussion	Modern taxonomic arrangements of the colobus monkeys either divide the red colobus and the Olive Colobus into two genera, Piliocolobus and Procolobus, respectively, or consider them to belong to one genus, Procolobus, with two subgenera (Procolobus for the Olive Colobus and Piliocolobus for the red colobus). We follow here C. P. Groves in his publications of 2001 and 2007 in using two genera. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF91FF95FAC5693BFDB7F755.taxon	distribution	Distribution. SE Nigeria (Ikpan block in the Oban Hills Division in Cross River National Park) and NW Cameroon (Yabassi, Korup National Park, Ebo Forest just N of the Sanaga River and Makombe Forest to the N of Ebo Forest, where they were heard in 2003).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF91FF95FAC5693BFDB7F755.taxon	description	Descriptive notes. Head-body 56 - 63 cm (males) and ¢. 62 cm (females), tail 75 — 76 cm (males) and ¢. 75 cm (females); no specific data are available for body weight. Fur of Preuss’s Red Colobus is dense and frizzier than in other species of Piliocolobus. Each body hair is ticked red and black, with black predominating on the dorsum and red on flanks. Middle of the back is dark brown, and lower flanks, tail, and limbs (including hands and feet) are bright orange-rufous. Underparts are pale red-gold; this color extends very narrowly up the throat to the chin. Inner surfaces of limbs are white. Cheeks are orange, forehead black, and crown and neck are blackish. Thereis a whorl above the brows, but no whorls above the ears. Infants are black, with gray underparts. Adult females have a large, prominentclitoris. In N. Ting’s mtDNA phylogeny, Preuss’s Red Colobus forms a clade with, and is very closely related to, Pennant’s Red Colobus (P. pennantiz), but its vocalizations are distinct.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF91FF95FAC5693BFDB7F755.taxon	biology_ecology	Habitat. Tall primary, lowland moist rainforest. Preuss’s Red Colobus is not found in secondary, regenerating forest.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF91FF95FAC5693BFDB7F755.taxon	food_feeding	Food and Feeding. Young leaves compose the vast majority of the diet of Preuss’s Red Colobus, supplemented by flowers and buds.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF91FF95FAC5693BFDB7F755.taxon	breeding	Breeding. Female Preuss’s Red Colobus display an enormous pink sexual swelling during the periovulatory period (the largest of all the red colobus), affecting both the anus and circumanal skin.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF91FF95FAC5693BFDB7F755.taxon	activity	Activity patterns. Preuss’s Red Colobusis diurnal and arboreal.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF91FF95FAC5693BFDB7F755.taxon	biology_ecology	Movements, Home range and Social organization. Very little is known about these aspects in Preuss’s Red Colobus. Home range size has been estimated at ¢. 100 ha. Group sizes of 20 to more than 80 individuals have been recorded.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF91FF95FAC5693BFDB7F755.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Critically Endangered on The IUCN Red List (as Procolobus preussi). Preuss’s Red Colobusis listed as Class B in the African Convention on the Conservation of Nature and Natural Resources. Major threats are hunting and habitat degradation. Hunting is particularly intense, making up a relatively large proportion of the total primate harvest around Korup National Park, especially in the north-east. In 1996, it was estimated that 10,000 - 15,000 Preuss’s Red Colobus may have been present in Korup, the stronghold for the taxon, but another estimate around that same time put the total population atjust 8000 individuals. In 1992, group densities in north-eastern Korup were estimated at 0 - 52 - 0 - 56 groups / km? which are not too dissimilar from more recent estimates of 0 - 46 groups / km? in 2008 in the same area. No absolute density is available for southern Korup, but J. Linder in 2008 encountered twelve groups in 243 km that he walked. It occurs in Korup and Ebo national parks in Cameroon and Cross River National Park in Nigeria. Recent surveys suggest it has been extirpated from Ejagham Forest Reserve in Cameroon.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF90FF95FF256DE4F725F866.taxon	materials_examined	Kenya, Tana River, Muniuni. (2 ° 6 ’ 24 ” S, 40 ° 6 ’ E).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF90FF95FF256DE4F725F866.taxon	discussion	Modern taxonomic arrangements of the colobus monkeys either divide the red colobus and the Olive Colobus into two genera, Piliocolobus and Procolobus, respectively, or consider them to belong to one genus, Procolobus, with two subgenera (Procolobus for the Olive Colobus and Piliocolobus for the red colobus). We follow here C. P. Groves in his publications of 2001 and 2007 in using two genera. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF90FF95FF256DE4F725F866.taxon	distribution	Distribution. SE coastal Kenya, confined to a narrow stretch of forest along the lower Tana River, mainly from Garsen N nearly as far as Wenje; recently also found in the forests of the Tana River Delta.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF90FF95FF256DE4F725F866.taxon	description	Descriptive notes. Head-body ¢. 50 cm, tail ¢. 65 cm; weight ¢. 9 kg. The Tana River Red Colobusis a very small species, nearly as small as the Zanzibar Red Colobus (P. kirkii). Males are larger than females, with a somewhat shorter tail. Skull is distinct, and strongly different from all other species of red colobus. It is very broad across orbits but narrow across canines, and teeth are notably large. Pelage is dark gray, with a slight buffy tinge above and a grayish-white underside. Limbs are lighter in color than the body, and tail darker, becoming nearly black toward the tip. Hands and feet are a little, if at all, darker than the rest of the limbs. Crown is brick-red and surrounded by a dark gray rim, with a whorl behind each ear leading to a low transverse crown crest. There is a whorl on either side of the crown above the ears, and the forehead and hairs over the ears are rimmed with black. Adult females have a large, prominent clitoris. DNA analysis places the Tana River Red Colobus close to other East African species, including the Ashy Red Colobus (P. tephrosceles), Lang’s Red Colobus (P. langi), and Foa’s Red Colobus (P. foa).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF90FF95FF256DE4F725F866.taxon	biology_ecology	Habitat. Relict evergreen gallery forest patches and narrow riparian forest strips, bordering rivers, streams, lakes, and wetlands in a landscape that otherwise has few trees.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF90FF95FF256DE4F725F866.taxon	food_feeding	Food and Feeding. The Tana River Red Colobus eats mainly young leaves, although flowers and fruit can make up a significant part ofits diet.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF90FF95FF256DE4F725F866.taxon	breeding	Breeding. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF90FF95FF256DE4F725F866.taxon	activity	Activity patterns. The Tana River Red Colobus is diurnal and arboreal. Studies have shown that it tends to spend 48 - 55 % ofits day resting, 23 - 30 % feeding, 7 - 24 % moving, and 2 - 7 % engaging in social activities.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF90FF95FF256DE4F725F866.taxon	biology_ecology	Movements, Home range and Social organization. Average daily movements of Tana River Red Colobus are 461 - 603 m, with home ranges of 9 - 13 ha. Group sizes are 4 - 24 individuals, with 1 - 2 adult males in each group. Dispersal is much more common in females than males. Reported densities are 33 - 253 ind / km?.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF90FF95FF256DE4F725F866.taxon	conservation	Status and Conservation. CITES Appendix I. Classified as Least Concern on The IUCN Red List (as Procolobus rufomitratus). The Tana River Red Colobus is listed as Class B in the African Convention on the Conservation of Nature and Natural Resources. It is threatened by changes in vegetation due to dam construction, irrigation projects, and water diversion that have altered the water table; forest clearing for agriculture; fires eroding levee forests; degradation due to livestock and firewood collection; selective felling of fig trees (Ficus, Moraceae) for canoes; and hunting. The area inhabited (25 km?) is broken into more than 70 patches, but two-thirds of the population occurs in the continuous forest belt along either side of the Tana River between Munazini and Makere ya Gwano, near the northern end of its distribution. The total population is estimated at 1100 - 1300 individuals, down from an estimated 1200 - 1800 in 1975. This is not considered a significant decrease and suggests that the Tana River Red Colobus may have developed strategies to cope with its shrinking habitat. Recent surveys indicate that at least 86 groups occur in 34 forest patches; mean group size has declined by c. 50 % since the 1970 s. It is now protected somewhat within the confines of the Tana River National Primate Reserve. Regardless, it has a very small extent of occurrence (52 km?), which is gradually becoming smaller and more fragmented. The Tana River Red Colobus occurs at less than five locations, and there is continuing decline due to the current rapid destruction of habitat. About 40 % of the remaining population is in 13 km? offorest in the 169 km * Tana River National Primate Reserve. Stringent habitat protection is required, and there is a need to reestablish the Mchelo Research Station. The proposed Tana Integrated Sugar Project in Tana River and Lamu districts threatens more than 200 km? of semi-natural habitat in the area. Tana River Red Colobus requires regular monitoring because it could very quickly become critically endangered.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF90FF96FA2E62F7FB70FDA5.taxon	materials_examined	Democratic Republic of Congo, Tungula (= Kilia), right bank of Semliki River.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF90FF96FA2E62F7FB70FDA5.taxon	discussion	Modern taxonomic arrangements of the colobus monkeys either divide the red colobus and the Olive Colobus into two genera, Piliocolobus and Procolobus, respectively, or consider them to belong to one genus, Procolobus, with two subgenera (Procolobus for the Olive Colobus and Pil 1 ocolobus for the red colobus). We follow here C. P. Groves in his publications of 2001 and 2007 in using two genera. Over a wide area in north-eastern DR Congo (broadly speaking, the Ituri Forest: from the mountains west of Lake Edward west nearly to the Lualaba, and from the Aruwimi-Ituri south to the River Maiko) occurs what is almost certainly a hybrid swarm between P. semlikiensis, P. oustaleti, and P. langi. Hybrids most frequently have bright red anterior body, shoulders, and arms; a dark “ orangey-brown ” mid-region; black rump, tail, and legs; and gray to yellow underparts, but they may be chestnut-brown, or they may lack the tripartite distribution of colors on their upperparts, or they may be mostly dark red-brown. The name P. ellioti (or it may be treated as if it were a subspecies of one of the other taxa) has been applied to these Ituri Forest populations, but there seemslittle doubt of their hybrid status. Red colobus occur in the Kahuzi-Biéga National Park, but their taxonomic status is unclear. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF90FF96FA2E62F7FB70FDA5.taxon	distribution	Distribution. NE DR Congo, in a very small area (c. 0 ° 20 ° - 0 ° 50 ’ N) between the middle Semliki River Valley and the Ruwenzori Mts (where there are no red colobus) to the E.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF90FF96FA2E62F7FB70FDA5.taxon	description	Descriptive notes. There are no specific measurements available. Body of the Semliki Red Colobusis anteriorly gray both above and below, with a blackish mantle. Arms are mostly brick-red, and hands are black. Legs are blackish-gray, with a trace of reddish, and tail is black. The black forehead band reaches the ears; a frontal tuft is present; throat, whiskers, and sides of head are reddish; and the nape is brown. There is a prominent tuft at the base of each ear, and a small “ panache ” (tuft of hair) is present.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF90FF96FA2E62F7FB70FDA5.taxon	biology_ecology	Habitat. Ironwood (Cynometra, Fabaceae) forest.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF90FF96FA2E62F7FB70FDA5.taxon	food_feeding	Food and Feeding. Diets include young leaves and mature leaves, fruit, flowers, buds, and possibly seeds.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF90FF96FA2E62F7FB70FDA5.taxon	breeding	Breeding. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF90FF96FA2E62F7FB70FDA5.taxon	activity	Activity patterns. The Semliki Red Colobus is diurnal and arboreal.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF90FF96FA2E62F7FB70FDA5.taxon	biology_ecology	Movements, Home range and Social organization. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF90FF96FA2E62F7FB70FDA5.taxon	conservation	Status and Conservation. CITES Appendix II. The Semliki Red Colobus has not been assessed on The IUCN Red List. No information is available concerning its conservation status.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF93FF96FFC667B2FB84F57E.taxon	materials_examined	Democratic Republic of Congo, Ouroua Mountains, between south-west shore of Lake Tanganyika and upper Congo-Lualaba.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF93FF96FFC667B2FB84F57E.taxon	discussion	Modern taxonomic arrangements of the colobus monkeys either divide the red colobus and the Olive Colobus into two genera, Piliocolobus and Procolobus, respectively, or consider them to belong to one genus, Procolobus, with two subgenera (Procolobus for the Olive Colobus and Piliocolobus for the red colobus). We follow here C. P. Groves in his publications of 2001 and 2007 in using two genera. Two subspecies may perhaps be recognized: foa: in the mountains along the western side of Lake Tanganyika and lulindicus in the lowland forest east of the Lualaba River. Considered monotypic here.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF93FF96FFC667B2FB84F57E.taxon	distribution	Distribution. EC DR Congo, lowlands E of the Lualaba River from the Lowa and Oso rivers in the N to the Elila River in the S, and Mts between the Lowa and Oso rivers in the N to ¢. 6 ° Sin the S, along the W side of Lake Tanganyika.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF93FF96FFC667B2FB84F57E.taxon	description	Descriptive notes. Head — body 50 - 69 cm (males), tail 62 - 67 cm (males); weight 9 - 13 kg (males) and 7 - 9 kg (females). Foa’s Red Colobus has long fur and a relatively short tail. Furis essentially red above, with a bit of black on the proximal part of the dorsum (although it may also be black, red, or black and red). Underparts are smoky-gray to yellowish-white. There is a crest on the forehead or anterior crown, and a prominent tuft at the base of each ear. Crown including the crest is red, contrasting with a black brow band and light colored cheeks. Limbs are wholly red, and hands and feet are generally dark to black. There is a prominent tuft of hair, or “ panache, ” on either side of the base ofthe tail. Teeth are small compared with most other species of red colobus. All Central African taxa of red colobus (excluding those south ofthe Grand Cuvette) are all curiously variable and contrast greatly with all other taxa that are noticeably homogeneous. It has been opined that Foa’s Red Colobus consists of two different taxa (one highland and one lowland) that have fused, and, if so, the components cannot now readily be disentangled, although there are average differences between highland and lowland populations and they may be recognized as subspecies. Analysis of mtDNA places Foa’s Red Colobus alongside mainly other species from East and Central Africa.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF93FF96FFC667B2FB84F57E.taxon	biology_ecology	Habitat. [. owland tropical rainforest in the west (on the right bank of the LLualaba) and montane rainforest in the east (Itombwe Mountains).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF93FF96FFC667B2FB84F57E.taxon	food_feeding	Food and Feeding. Diets include young leaves and mature leaves, fruit, flowers, buds, and possibly seeds.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF93FF96FFC667B2FB84F57E.taxon	breeding	Breeding. Female Foa’s Red Colobus displays a substantial pink sexual swelling during the periovulatory period.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF93FF96FFC667B2FB84F57E.taxon	activity	Activity patterns. Foa’s Red Colobusis diurnal and arboreal	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF93FF96FFC667B2FB84F57E.taxon	biology_ecology	Movements, Home range and Social organization. There is no information available for this species in the wild.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF93FF96FFC667B2FB84F57E.taxon	conservation	Status and Conservation. CITES Appendix II. Foa’s Red Colobus has not been assessed on The IUCN Red List. It is heavily hunted for the bushmeat trade, but otherwise no information is available concerning its conservation status.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF93FF96FFFA6FDAF724FBF5.taxon	materials_examined	Democratic Republic of Congo, Risimu, between Stanleyville (= Kisangani) and Bafwaboli.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF93FF96FFFA6FDAF724FBF5.taxon	discussion	Modern taxonomic arrangements of the colobus monkeys either divide the red colobus and the Olive Colobus into two genera, Piliocolobus and Procolobus, respectively, or consider them to belong to one genus, Procolobus, with two subgenera (Procolobus for the Olive Colobus and Piliocolobus for the red colobus). We follow here C. P. Groves in his publications of 2001 and 2007 in using two genera. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF93FF96FFFA6FDAF724FBF5.taxon	distribution	Distribution. NE DR Congo, restricted to the “ cul-de-sac ” between the Lualaba and Aruwimi-Ituri rivers.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF93FF96FFFA6FDAF724FBF5.taxon	description	Descriptive notes. Head-body ¢. 50 cm (males) and 48 - 5 cm (females), tail 66 - 5 cm (males) and 44 - 66.5 cm (females); weight ¢. 9 kg (males) and c. 7 - 7 kg (females). Head, throat, chest, arms, and shoulders of Lang’s Red Colobus are reddish, contrasting with the sepia-black tone of hindparts and belly. Individuals resembling this description occur throughout the distribution of the “ elliot ” hybrid swarm (described under the Semliki Red Colobus, P. semlikiensis). The mtDNA affinities of Lang’s Red Colobus are, as one would expect, with its neighbors in north-eastern DR Congo and with P. tephrosceles.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF93FF96FFFA6FDAF724FBF5.taxon	biology_ecology	Habitat. Lowland rainforest.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF93FF96FFFA6FDAF724FBF5.taxon	food_feeding	Food and Feeding. Diets include young leaves and mature leaves, fruit, flowers, buds, and possibly seeds.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF93FF96FFFA6FDAF724FBF5.taxon	breeding	Breeding. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF93FF96FFFA6FDAF724FBF5.taxon	activity	Activity patterns. Lang’s Red Colobus is diurnal and arboreal.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF93FF96FFFA6FDAF724FBF5.taxon	biology_ecology	Movements, Home range and Social organization. There is no information available for this species in the wild.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF93FF96FFFA6FDAF724FBF5.taxon	conservation	Status and Conservation. CITES Appendix II. Lang’s Red Colobus has not been assessed on The IUCN Red Lust. It is not found in any protected areas, and no information is available concerning its conservation status.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF93FF97FAC66145FC5BFD85.taxon	materials_examined	Youmba, right bank of lower Oubangui (= Ubangi), 0 ° 18 ’ N, 17 ° 30 ’ E.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF93FF97FAC66145FC5BFD85.taxon	discussion	Modern taxonomic arrangements of the colobus monkeys either divide the red colobus and the Olive Colobus into two genera, Piliocolobus and Procolobus, respectively, or consider them to belong to one genus, Procolobus, with two subgenera (Procolobus for the Olive Colobus and Pil- 1 ocolobus for the red colobus). We follow here C. P. Grovesin his publications of 2001 and 2007 in using two genera. The southwestern most locality, Pombo (1 ° 13 ° 127 S, 16 ° 9 ° 36 ” E), is very close to Butando where P. bouvieri is found, but there is no indication of intermediates between them. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF93FF97FAC66145FC5BFD85.taxon	distribution	Distribution. Extreme S South Sudan, S Central African Republic, N DR Congo (N to Sili, Uele District, 4 ° 2 ° 24 ” N, 27 ° 6 ’ 36 ” E, Dedegwa, Garamba, 4 ° 20 ° N, 29 ° 25 ’ 48 ” E, S to the Congo River, and E to Lake Albert), NE Republic of the Congo (N to Mongoumba, 3 ° 23 ’ N, 18 ° 21 ' E, and Bakota, S bank of the Lobaye River, 3 ° 22 " 43 ” N, 17 ° 20 ° 24 ” E, and SW to Pombo, Sangha Department).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF93FF97FAC66145FC5BFD85.taxon	description	Descriptive notes. Head-body 45.9 - 68 cm (males) and 52 - 64 cm (females), tail 55 - 5 — 73 cm (males) and 68 - 73 cm (females); weight 12 - 5 kg (males) and 8 - 2 kg (females). Oustalet’s Red Colobus is a large, robust, sexually dimorphic species, with a short tail. It is dark speckled smoky-brown (sometimes brownish-fawn) above, and slightly paler below (buffy to whitish or pale red). The crown does not, or hardly, contrasts with back. The whiskers are grayish. Hands and feet are dark to black. This is another very variable Central African species. Some individuals in the Ubangi / Lobaye area are described as “ raw sienna brown, ” underparts may be golden, and tail root has a coppery red patch. Those from the Uele River (north-eastern DR Congo) gallery forest region tend to be lighter brownish-fawn, with forelimbs and lower legs especially pale, contrasting with dark hands and feet. Finally, there is a deep red-colored morph both in the south-western part of the distribution and in the extreme east, on the borders of the Ituri Forest, the latter probably due to gene-flow from the hybrid swarm described under the Semliki Red Colobus (P semlikiensis). In both mtDNA and vocalizations, Oustalet’s Red Colobus groups with other species of red colobus in East Central Africa. Nevertheless, like other species of red colobus in the Congo Basin, Oustalet’s Red Colobus carries at least two very divergent lineages of mtDNA, with different specimens from different sites clustering with other red colobus species instead of with one another. A prominent tuft of hair (“ panache ”) is present, although it is shorter than on the Tshuapa Red Colobus (P. tholloni). Like Foa’s Red Colobus (P. foai), Oustalet’s Red Colobus has small teeth. Cranially, Oustalet’s Red Colobus, Foa’s Red Colobus, the Semliki Red Colobus, and the Ashy Red Colobus (P. tephrosceles) form a group — all have a long palate, early closing skull sutures, and angular orbits.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF93FF97FAC66145FC5BFD85.taxon	biology_ecology	Habitat. Mature primary lowland, tropical moist forest, swamp and gallery forest, and savanna woodland. In the Ngotto Forest of the Central African Republic, however, Oustalet’s Red Colobus only occurs in flooded forests on alluvial riverbanks.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF93FF97FAC66145FC5BFD85.taxon	food_feeding	Food and Feeding. Diets include young leaves and mature leaves, fruit, flowers, buds, and possibly seeds. One study reported that groups regularly entered water to collect bulbs of aquatic plants.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF93FF97FAC66145FC5BFD85.taxon	breeding	Breeding. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF93FF97FAC66145FC5BFD85.taxon	activity	Activity patterns. Oustalet’s Red Colobusis diurnal and arboreal.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF93FF97FAC66145FC5BFD85.taxon	biology_ecology	Movements, Home range and Social organization. Group sizes of Oustalet’s Red Colobus were 3 - 18 individuals at one site. Groups there spent 30 % of their time in the lower forest canopy, less than 10 m above the ground.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF93FF97FAC66145FC5BFD85.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Least Concern on The IUCN Red List (as Procolobus rufomitratus oustaleti). Oustalet’s Red Colobus is likely to be among the more numerous of the red colobus taxa, given its very wide distribution north of the Congo River where there are still large areas of forest and low human densities. Nevertheless, it is heavily hunted for the bushmeat trade. It occurs in Okapi Wildlife Reserve and Tayna Gorilla Reserve in DR Congo and Nouabalé-Ndoki National Park in the Republic of the Congo. It is also found in Ngotto Classified Forest and the proposed Mbaéré-Bodingué National Park in the Central African Republic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF92FF97FF3767D1FAAEF6C3.taxon	materials_examined	Democratic Republic of Congo, Mabobi (0 ° 0 ’ 36 ” N, 25 ° 11 ° 24 ” E), left bank of Lualaba (= Upper Congo) River.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF92FF97FF3767D1FAAEF6C3.taxon	discussion	Modern taxonomic arrangements of the colobus monkeys either divide the red colobus and the Olive Colobus into two genera, Piliocolobus and Procolobus, respectively, or consider them to belong to one genus, Procolobus, with two subgenera (Procolobus for the Olive Colobus and Piliocolobus for the red colobus). We follow here C. P. Groves in his publications of 2001 and 2007 in using two genera. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF92FF97FF3767D1FAAEF6C3.taxon	distribution	Distribution. C DR Congo, between the Lomami and Lualaba rivers, S to about the Ruiki and Lutanga rivers.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF92FF97FF3767D1FAAEF6C3.taxon	description	Descriptive notes. There are no specific measurements available; weight 9.9 - 2 kg (males) and 7 - 5 kg (females). The Lomami Red Colobus is among the smallest of the Central African red colobus, and it has a longer tail than the Tshuapa Red Colobus (P. tholloni). Furis red with black on the mid-back, shoulders, and upper arms. Underside, and inner surfaces of limbs are grayish-white. Hands and feet are contrastingly black. Tail is reddish basally, becoming gradually darker along its length but never quite black. Crown is speckled reddish, bordered in front by a black band that extends to the temples. A long black tuft is present at the base of each ear. Facial skin is black, except for narrow depigmented zones along lips, chin, base of the nose, and upper eyelids. The Lomami Red Colobus is unusual among Central African red colobus in having no prominent tuft on the head (“ panache ”). Its mtDNA affinities are with other East Central African red colobus species, but it does not seem particularly closely related to its neighbors in eastern DR Congo.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF92FF97FF3767D1FAAEF6C3.taxon	biology_ecology	Habitat. Mostly lowland tropical rainforest.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF92FF97FF3767D1FAAEF6C3.taxon	food_feeding	Food and Feeding. Diets include young leaves and mature leaves, fruit, flowers, buds, and possibly seeds.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF92FF97FF3767D1FAAEF6C3.taxon	breeding	Breeding. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF92FF97FF3767D1FAAEF6C3.taxon	activity	Activity patterns. Lomami Red Colobus is diurnal and arboreal.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF92FF97FF3767D1FAAEF6C3.taxon	biology_ecology	Movements, Home range and Social organization. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF92FF97FF3767D1FAAEF6C3.taxon	conservation	Status and Conservation. CITES Appendix II. The Lomami Red Colobus has not been assessed on The IUCN Red List. It is not found in any protected areas, and no information is available concerning its conservation status.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF92FF97FF346C97FA1AF473.taxon	materials_examined	Uganda, Toro, Ruahara River, east side of Mount Rwenzori, 1200 m.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF92FF97FF346C97FA1AF473.taxon	discussion	Modern taxonomic arrangements of the colobus monkeys either divide the red colobus and the Olive Colobus into two genera, Piliocolobus and Procolobus, respec-: tively, or consider them to belong to one genus, Procolobus, with two subgenera (Procolobus for the Olive Colobus and Piliocolobus for the red colobus). We follow here C. P. Grovesin his publications of 2001 and 2007 in using two genera. P. tephrosceles was considered to be the eastern Rift Valley subspecies of P. oustaleti by J. Kingdon. There is some geographic variation in this species, although further study is needed to determine if they merit subspecific rank. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF92FF97FF346C97FA1AF473.taxon	distribution	Distribution. E border of the W Rift Valley from c. 1 ° N to 8 ° S in isolated forests in SW Uganda (Kibale), and W Tanzania (Biharamulo, Gombe, Mahale Mts, and Lake Rukwa); possibly in E DR Congo, Rwanda, and Burundi, butits presence has not yet been confirmed.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF92FF97FF346C97FA1AF473.taxon	description	Descriptive notes. Head-body 45.5 - 67 cm (males) and 49 - 58 cm (females), tail 65 — 72 cm (males) and 62 - 68 cm (females); weight 8 - 11.4 kg (males); no weight data available for females, but they are considerably smaller than males. The Ashy Red Colobus has long, dense fur and a relatively short tail. It is extremely sexually dimorphic. Males are considerably heavier than females, with a more massive head, larger canines, and longer cheek whiskers. A small, light-colored “ panache ” (a prominent tuft of hair) is present. It is glossy-black above (including tail) and light gray to white below. Forearms and legs are a dull light gray, occasionally with varying amounts of tan or brown on arms, and often redder tones on haunches and upper thighs. Crown is rusty-red with a marked, dark reddish-brown forehead crest bordered by a black stripe on the temples. Face is slaty (except for a trace of pale pinkish tones on eyelids) and outlined with gray whiskers. There is a prominenttuft at the base of each ear. Callosities are slaty or pink. Adult females have a large, prominent clitoris. Its mtDNA affinities are with other East Central African red colobus species, particularly Lang’s Red Colobus (P. langi) and eastern QOustalet’s Red Colobus (PF. oustaleti). There is some geographic variation in color, and it may be that subspecies could be recognized. Populations in the Mahale Mountains (western Tanzania, bordering Lake Tanganyika), for example, tend to be jet-black with restricted red areas on hindquarters, tail becomes black distally, and underside is white. In forests around Lake Rukwa, the southernmost part of its range, populations are more blackish-gray, underside is light gray, sideburns are red, and cheek whiskers are very long. Teeth are relatively small, and the skull is strongly sexually dimorphic in size and is instantly recognized by a curious furrow or crease running between the orbits across the base of the nasal bones.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF92FF97FF346C97FA1AF473.taxon	biology_ecology	Habitat. Lowland and montane primary rainforest. In the Kibale Forest of Uganda, the Ashy Red Colobus is most abundant in moist, old growth, evergreen forest, with emergent trees reaching 50 m in height. It can persist in lightly logged forest if important food trees remain.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF92FF97FF346C97FA1AF473.taxon	food_feeding	Food and Feeding. [. eaves make up the vast majority of the diet of the Ashy Red Colobus, although the proportion of young leaves to mature leaves varies between sites and seasons. Flowers and fruit make up a small component of the diet.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF92FF97FF346C97FA1AF473.taxon	breeding	Breeding. The female Ashy Red Colobus displays a small pink sexual swelling during the periovulatory period. Breeding does not appear to be seasonal, and the interbirth interval is 25 - 5 months. Neonates have black silky fur on their back and sides and a gray ventral surface; muzzle, ears, palms, and soles are pink. Subadult males have a relatively small, knob-like perineal organ.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF92FF97FF346C97FA1AF473.taxon	activity	Activity patterns. The Ashy Red Colobus is diurnal and arboreal. At Gombe, Tanzania, T. Clutton-Brock found that it spent 54 % of the day resting, 25 % feeding, 9 % engaging in social activities, 8 % moving, and 4 % in various other activities. T. Struhsaker, studying the Ashy Red Colobus in Kibale, Uganda, found somewhat different activity patterns: 45 % feeding, 38 % resting, 9 % moving, and 8 % engaging in social activities.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF92FF97FF346C97FA1AF473.taxon	biology_ecology	Movements, Home range and Social organization. Mean daily movement is 649 m in a 65 ha home range. Group sizes are 9 - 80 individuals, with multiple adult males and females. Females are likely to transfer from their natal groups more often than males.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF92FF97FF346C97FA1AF473.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Endangered on The IUCN Red Last (as Procolobus rufomitratus tephrosceles). The main threat to the Ashy Red Colobus is habitat loss through selective logging of mature forest for timber, conversion to agricultural land, production of charcoal, and forest fires. To some degree, hunting for meat and skins remains a threat. The Ashy Red Colobus numbers at least 20,000 individuals, with the largest and most viable population in Kibale Forest in Uganda where the population is estimated at 17,000 or more. About 60 % of the Kibale Forest National Park has suitable habitat for red colobus, and published density estimates in this habitat are 93 - 313 ind / km ®. Although Kibale would appear to have a healthy and viable population, long-term studies in two areas of the Park indicate statistically significant declines in numbers of Ashy Red Colobus. In the Ngogo study area, census data spanning nearly 24 years indicated a 43 % decline in groups, mainly due to predation by Chimpanzees (Pan troglodytes). The Ashy Red Colobus in Gombe Stream National Park also seem to have declined in numbers due to predation by Chimpanzees. Red colobus group sizes have declined by nearly 50 % over a 25 year period and 16 - 40 % of them are killed each year by Chimpanzees. This is particularly troubling given the very small size (¢. 80 km?) and isolation of the Gombe Stream National Park. Besides the Gombe Stream and Kibale Forest national parks, the Ashy Red Colobusis found in Birigi Game Reserve and Mahale Mountains National Park in Tanzania. The population in Kibale Forest National Park may be the only viable one, although recent survey work by T. Davenport and colleagues has shown that at least 1354 Ashy Red Colobus survive on the Ufipa Plateau, with 1217 individuals in Mbizi and 137 individuals in Mbuzi. Long-term monitoring of the Ngogo situation is required, as well as a comprehensive, long-term monitoring program for the colobus in Kibale.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF92FF90FA316EC7FBA0F86E.taxon	materials_examined	Democratic Republic of Congo, Lower Congo.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF92FF90FA316EC7FBA0F86E.taxon	discussion	Modern taxonomic arrangements of the colobus monkeys either divide the red colobus and the Olive Colobus into two genera, Piliocolobus and Procolobus, respectively, or consider them to belong to one genus, Procolobus, with two subgenera (Procolobus for the Olive Colobus and Piliocolobus for the red colobus). We follow here C. P. Groves in his publications of 2001 and 2007 in using two genera. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF92FF90FA316EC7FBA0F86E.taxon	distribution	Distribution. DR Congo E of the Congo River and S of the great bend of the river, the E limit is not the Lualaba Riveritself, but rather the Lomami River, and bounded by the Kasai and Sankuru rivers in the S.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF92FF90FA316EC7FBA0F86E.taxon	description	Descriptive notes. Head-body c. 64 cm (males) and 58 - 60 cm (females), tail c. 63 cm (males) and 41 - 69 cm (females); no specific data are available for body weight. On the basis of only three specimens, the Tshuapa Red Colobus seems to be a rather large species, with a very short tail. Fur is uniformly foxy-red above, although somewhat darker on shoulders and tail, and creamy-white below. Crown is light bay to mahogany. Circumfacial hair is short and black, with only posterior parts of cheeks light-colored like underparts. Legs are orange-rufous, paler than arms, and hands, feet, and tail tip are dark to black. There are long black tufts at the base ofthe tail. Hairs at the sides ofthe tail-base develop into along “ panache ” (tuft). Face is slaty, with a trace of pale on eyelids and lips. Ischial callosities are pink or slaty. Like other East Central African red colobus, mtDNA analysis shows unexpected heterogeneity; specimens from the same locality show very divergent mitochondrial lineages. Skull of the Tshuapa Red Colobus is distinctively long and narrow, small-toothed, and very prognathous, with a large sagittal crest, which is more extensive than in other forms.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF92FF90FA316EC7FBA0F86E.taxon	biology_ecology	Habitat. Lowland flooded moist primary rainforest.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF92FF90FA316EC7FBA0F86E.taxon	food_feeding	Food and Feeding. The Tshuapa Red Colobus eats mostly young leaves, but seeds can be a substantial part of its diet.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF92FF90FA316EC7FBA0F86E.taxon	breeding	Breeding. The female Tshuapa Red Colobus displays a substantial pink sexual swelling during the periovulatory period.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF92FF90FA316EC7FBA0F86E.taxon	activity	Activity patterns. The Tshuapa Red Colobusis diurnal and arboreal.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF92FF90FA316EC7FBA0F86E.taxon	biology_ecology	Movements, Home range and Social organization. The Tshuapa Red Colobus has never been studied in the wild and very little is known about it. Groups of more than 60 individuals have been reported.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF92FF90FA316EC7FBA0F86E.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Near Threatened on The IUCN Red Lust (as Procolobus rufomitratus tholloni). The Tshuapa Red Colobusis believed to have undergone a population decline on the order of 20 - 25 % over the last three generations, due mainly to intensive hunting (particularly during the period of the Congo War, 1996 - 2003). Habitat loss for timber and agricultural land likely affects some populations locally. Further research into hunting may reveal that the Tshuapa Red Colobus could qualify as Vulnerable, especially because it is more heavily hunted than many of the other primates in the Congo Basin. It occurs in Salonga National Park and Tayna Gorilla Reserve in DR Congo.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF95FF90FFE66DB5F932F41D.taxon	materials_examined	Africa.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF95FF90FFE66DB5F932F41D.taxon	discussion	This species is monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF95FF90FFE66DB5F932F41D.taxon	distribution	Distribution. West Africa, from SE Sierra Leone and S Guinea to Ghana (extending just E of the Volta River), and in S Togo, S Benin, and S Nigeria (S bank of the River Benue just above its confluence with the Niger River in the Idah, Kogi State, and the lower Niger). There is a discontinuity in the distribution between the VoltaRiver in Ghana and the Nigerian population, and it is believed that its range was far greater in the recent past.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF95FF90FFE66DB5F932F41D.taxon	description	Descriptive notes. Head-body 46 - 51 cm (males) and 38 - 55 cm (females), tail 50 - 61 cm (males) and 43 - 71 cm (females); weight 4.5 - 7 kg (males) and 3.4 - 2 kg (females). The Olive Colobus is the smallest of the colobine monkeys. On the basis of hunter-shot specimens and museum records, J. Oates and colleagues gave the following body weights: males 4 - 7 kg (4.5 - 7 kg; n = 20) and females 4 - 2 kg (2.8 - 5.4 kg; n = 14). The Olive Colobus is generally a dull olive-brown dorsally, becoming grayish-white below and on limbs. Head is small and rounded, with a virtually hairless face and an off-white circumfacial ruff. There is a low longitudinal crest of upright hairs on the crown (more pronounced in the male), which is formed by a pair of lateral pale whorls. Tail is slender and untufted. Feet are enormous and can equal as much as onethird the length of the entire hindlimb. Thumbs are mere tubercles. Ischial callosities remain separate in both sexes.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF95FF90FFE66DB5F932F41D.taxon	biology_ecology	Habitat. Swamp forest, moist forest, dry semi-deciduous forest, dense regenerating forest, and abandoned farmland. The Olive Colobus prefers dense tangled growth in the lower levels of the forest canopy, and it is especially common in swampy riparian forest. It is rarely seen in undisturbed high forest and avoids feeding in the crowns of tall trees and going to the ground.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF95FF90FFE66DB5F932F41D.taxon	food_feeding	Food and Feeding. The diet of the Olive Colobus is composed mainly of young leaves from trees and climbing plants. Liana leaves are a year-round staple. Seeds and flowers are also important. In a 1988 study by J. Oates in Tiwai Island Wildlife Sanctuary, Sierra Leone, a group of Olive Colobus were highly selective feeders. The group ate from a small number of uncommon middle canopy trees, from the understory, and from climbers. Their food trees tended to be deciduous or produce large crops of new leaves over a short period of time. Unlike other colobus monkeys, mature leaves were largely ignored. They were eaten in small quantities only toward the end of the dry season when preferred items (young leaves and seeds) were least available. They obtained their year-round staple from young leaves; seeds, flowers, and leaf petioles were important seasonally. In the dry season, availabilities of seeds are higher and young leaves lower. Young leaves of Terminalia iworensis (Combretaceae), Sapium aubreville: (Euphorbiaceae; young leaves and mature leaf petioles), and Acacia pennata (Fabaceae) were predominant in the diet. Their preferred leaves tended to have low fiber and tannin levels. Overall 50 plant species were included in the diet. They did not seem to look for or eat small insects.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF95FF90FFE66DB5F932F41D.taxon	breeding	Breeding. Even before the advent of puberty, the female Olive Colobus begins to exhibit a substantial pink perineal swelling during the periovulatory period; a similar swelling is described for newborns. Females carry conspicuous sexual swellings for c. 50 % of their menstrual cycle, and this makes them conspicuous to males. Females are strongly proceptive and mate overwhelmingly with the dominant male in a group. Subadult males have a perineal organ. The Olive Colobus is, to some extent, a seasonal breeder, with a mating season in April-August and births in November-February after a gestation of 150 - 180 days. A single young is born. Female Olive Colobus are unique among Old World monkey mothers in that they carry their newborns dangling from their mouth for the first few weeks. Neonatal pelage resembles that of the adult, but it is slightly lighter and a bit more yellow.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF95FF90FFE66DB5F932F41D.taxon	activity	Activity patterns. The Olive Colobus is diurnal and almost exclusively arboreal. It is very shy. At Tiwai Island Wildlife Sanctuary, it spent 40 % of its day resting, 27 % feeding, 25 % moving, and 8 % engaging in social and other activities.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF95FF90FFE66DB5F932F41D.taxon	biology_ecology	Movements, Home range and Social organization. Average daily movement has been recorded at 1200 m, with home ranges of 28 - 58 ha. The Olive Colobuslives in small groups of 5 - 20 individuals (but usually under ten), with 1 - 2 adult males and the same number of adult females. Group membership is relatively fluid, with young individuals of both sexes dispersing from their natal groups, although adult females change groups more often than do adult males. The Olive Colobus is almost always found in close association with guenons (Mona, Cercopithecus mona, Spot-nosed, C. petaurista, Campbell's, C. campbelli, and Diana, C. diana, monkeys), and it responds to their alarm calls. These associations with other species are invariable, particularly with Diana Monkeys. Each Olive Colobus group follows “ its ” Diana Monkey group around, and territorial encounters between groups of Diana Monkeys bring their associated Olive Colobus into proximity, resulting in territorial counter-calling by adult males and extragroup matings.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF95FF90FFE66DB5F932F41D.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Near Threatened on The IUCN Red List. The Olive Colobusis listed as Class A in the African Convention on the Conservation of Nature and Natural Resources. It is probably declining in numbers because of widespread loss of habitat and hunting for food, especially in the far eastern part of its distribution toward Nigeria. Their cryptic nature and ability to survive in swamps and small forest fragments and to adapt to some anthropogenic disturbance have no doubt enabled them to persist and remain common in at least parts of their range. The Olive Colobus occurs in Bia, Kakum, and Nini-Suhien national parks, Bomfobiri Wildlife Sanctuary, Ankasa Game Reserve, and Sukusuku Reserve in Ghana; Tai Forest National Park in Ivory Coast; Mount Nimba Strict Nature Reserve and Sapo National Park in Liberia; and Gola Forest Reserves and Tiwai Island Wildlife Sanctuary in Sierra Leone.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF96FF93FF386710F9DCFC73.taxon	materials_examined	Indonesia, western Java.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF96FF93FF386710F9DCFC73.taxon	discussion	Genetically, both subspecies of P. comata are clearly distinct, but it has been shown that a chain of intermediate populations exists between them. D. Brandon-Jones has argued that fredericae warrants specific status. Two subspecies recognized.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF96FF93FF386710F9DCFC73.taxon	distribution	Subspecies and Distribution. P. c. comataDesmarest, 1822 — WJava, fromUjungKulnotoMtLawu; Elimitisunknown. P. c. fredericae Sody, 1930 — C Java, around the slopes of the volcanoes Gunung Slamet, Gunung Cupu, and Diyeng; the extent of its distribution to the W is not known.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF96FF93FF386710F9DCFC73.taxon	description	Descriptive notes. Head-body 48 - 60 cm (males) and 43 - 57 cm (females), tail 59 - 72 cm (males) and 56 - 72 cm (females); weight 6.7 - 8 kg (males) and 6 - 4 kg (females). The Javan Langur is dark gray or black above, with at least some white below. There is a single whorl of hair on front of crown, in addition to a short brow fringe, and the tall, miter-like crest behind is black, without stripes. The “ Javan Grizzled Langur ” (P. ¢. comata) is slaty-gray above and darker on limbs, crown, and dorsal side oftail. Underside, inner surfaces of arms and legs, and ventral surface of tail are whitish. Crown crest is black. Pelage on body is longer than on limbs or tail. The “ Javan Fuscous Langur ” (P. c. fredericae) is a melanistic form, black above and below, except for lower abdomen and inner surfaces of limbs, which are white, and throat and upper chest, which are light gray or whitish, the two zones being joined by a thin whitish streak. There is a small amount of white on digits.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF96FF93FF386710F9DCFC73.taxon	biology_ecology	Habitat. Primary and secondary rainforest, from lowlands to ¢. 2500 m, plantations, and orchards. The Javan Langur prefers the middle and upper layers of the forest canopy. It was historically found from sea level to 2565 m, but is now largely restricted to fragments of montane and upper montane forest. In Pegunungan Dieng, itis found in primary and secondary forests, at edges and in the interior, and in lowland forests, forests on steep slopes and hills, and upper montane forest. Rainfall in the interior ofJava can be very high; annual rainfall in the area of Mount Slamet is 5000 - 6714 mm / year.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF96FF93FF386710F9DCFC73.taxon	food_feeding	Food and Feeding. The diet of the Javan Langur consists mainly of young leaves, along with fruits, flowers, seeds, and fungi. Individuals supplement their diets with large tree snails, and also eat reddish soil. A study in Gede-Pangrango National Park recorded a diet of 63 % young leaves, 17 % flowers, 6 % mature leaves, 6 % fruits, and 2 % seeds (4 % unidentified). The study by Y. Ruhiyat recorded a similar diet: 59 % young leaves, 135 % fruits, and the remainder flowers, mature leaves, fungi, pseudo-bulbs, branch tips, and seeds. Preferred leaves include those of Ficus pubinervis (Moraceae), Passiflora ligularis (Passifloraceae), Elaeagnus triflora (Elacagnaceae), Schefflera aromatica (Araliaceae), Jasminum azoricum (Oleaceae), Hoya (Apocynaceae), and Aeschynanthus (Gesneriaceae). Preferred fruits include those of Premna parasitica (Lamniaceae), Saurauwia (Actinidiaeceae), and Castanopsis argentea (Fagaceae). Langurs in general are reported to avoid sweet succulent fruits such as figs, which can acidify the neutral pH of the forestomach and thus destroy beneficial gut microflora essential to leaf digestion.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF96FF93FF386710F9DCFC73.taxon	breeding	Breeding. There is no set breeding season of the Javan Langur. The female gives birth to a single young after a relatively short gestation period. Infants are carried ventrally until they are c. 1 year old. They are passed among the females in a group (allomothering) and are independent at c. 1 - 5 years old. Unlike other species of Presbytis, the natal coat of the Javan Langur is medium to dark gray or black.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF96FF93FF386710F9DCFC73.taxon	activity	Activity patterns. Javan Langurs are arboreal and diurnal. In Gede-Pangrango National Park, their activity budget was 51 % resting, 23 % foraging and feeding, 21 % traveling, and 5 % engaging in social and other activities. They have three to four feeding bouts during the day. They travel and feed in the upper canopy and sleep in high trees, generally on ridges.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF96FF93FF386710F9DCFC73.taxon	biology_ecology	Movements, Home range and Social organization. The Javan Langur lives in small groups, generally with a single adult male but sometimes with two. Surveys around Mount Slamet recorded groups of 2 - 14 individuals. They are quite territorial and encounters between groups are antagonistic. The adult male is dominant, and grooming is largely a female activity. Males disperse from their group before puberty. Sometimes they form mixed-species groups with the East Javan Langur (7 Trachypithecus auratus). Surveys of the Javan Fuscous Langur around Mount Slamet in central Java in 2006 and 2008 estimated 5 - 6 ind / km? on southern slopes to 8 ind / km? ® on the eastern slopes.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF96FF93FF386710F9DCFC73.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Endangered on The IUCN Red List, including both subspecies (under P. comata). The Javan Langur is severely threatened by habitat loss and, to a lesser extent, hunting. In 2006, the total population was estimated at 2300 - 2500 individuals in ¢. 30 isolated populations. No subpopulation contains more than 250 mature individuals. The Javan Grizzled Langur is threatened by habitat destruction, such that its remaining populations are now mainly confined to montane forest patches, whereas it once occurred across extensive lowland areas. Hunting is a persistent threat, especially in the area of Mount Slamet. Forests are gradually degraded through illegal logging, occasionalfelling for building materials, cutting trees to obtain orchids and pitcher plants, and the collection of firewood. Javan Langurs are also persecuted for crop raiding. They occur in at least five protected areas: Gede-Pangrango National Park, Gunung Halimun National Park, Gunung Tukung Gede Nature Reserve, Ujung Kulon National Park (Javan Grizzled Langur), and Gunung Slamet Nature Reserve (Javan Fuscous Langur).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF96FF9CFA0766C7FBA6FB32.taxon	materials_examined	Singapore.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF96FF9CFA0766C7FBA6FB32.taxon	discussion	P. femoralis is yet another highly variable langur, particularly in the color pattern of its head and in the direction of hairs on its chest. Until recently it was considered a subspecies of P. melalophos. Three subspecies recognized.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF96FF9CFA0766C7FBA6FB32.taxon	distribution	Subspecies and Distribution. P. f. femoralisMartin, 1838 — SPeninsularMalaysia (extremeSofPahangandJohorstates) andSingapore. P. f. percuraLyon, 1908 — CESumatra, inasmallareabetweentheRokanandSiakrivers. P. f. robinsoni Thomas, 1910 — S Myanmar, S Thailand (from 13 ° 50 ’ N in Phetchaburi Province), and NW Peninsular Malaysia (S to Larut Hills, Perak State).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF96FF9CFA0766C7FBA6FB32.taxon	description	Descriptive notes. Head-body 43 - 61 cm, tail 61 - 84 cm; weight 5.9 - 8.2 kg. The Banded Langur is generally dark brown or blackish above, with a white underside (continuous with a prominent zone of white on outsides of thighs). Interorbital region is wider than in the Black-crested Sumatran Langur (P. melalophos), and white eye-rings are present. Crown crest is shaggy and ill-defined. “ Raffles’s Banded Langur ™ (FP. f. femoralis) is blackish above with a red-brown head and a gray underside bisected by a white median line. There is no white under tail or on limbs. A pair of whorls is present on crown, with a long crest between (or, alternately, one of the pair may be suppressed). Hairs are directed backward on chest, forming a short crest where they meet forward-directed belly hairs. Singaporean individuals are darker ventrally and are slightly smaller than those from the mainland. “ Robinson’s Banded Langur ” (P. f. robinsoni) is gray-brown or blackish-brown, with a white underside. Chest, hands, and feet are black, and inner surfaces of limbs are white nearly to heels. Face (including eyelids) is black, with only a patch around the mouth being white. There are either one or two whorls on the crown. Whitish morphs have been reported. The “ East Sumatran Banded Langur ” (P. f. percura) is mainly blackish above, with a gray head and a white underside. White underside is narrow but extends horizontally across chest, down inner sides of limbs in thin stripes to wrists and ankles, and to cheeks and chin. Part of the ventral surface oftail is indistinctly white. Its throat and face are gray, and there are narrow, pale eyerings. One (sometimes two) whorls are set well back on its crown. Hairs on the chest are directed backward.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF96FF9CFA0766C7FBA6FB32.taxon	biology_ecology	Habitat. Primary and secondary rainforest, swamp, mangrove forest, and rubber tree plantations. The Banded Languris found in taller trees of swampy peat forest on the Malay Peninsula, but in Singapore, it is found in primary and secondary forest and swamp and dry-land rainforests.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF96FF9CFA0766C7FBA6FB32.taxon	food_feeding	Food and Feeding. The Banded Langur eats mainly young leaves and fruits, along with seeds, flowers (Myristica superba, Myristicaceae), buds, and fungi. In one study, fruits comprised more than 50 % of the diet. It also ingests soil.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF96FF9CFA0766C7FBA6FB32.taxon	breeding	Breeding. Female Banded Langurs usually initiate sexual interactions. A proceptive female approaches a male and crouches in front of him, presenting her hindquarters with her tail drooping for c. 5 seconds. She crouches and lifts her hindquarters repeatedly. The male usually raises her tail and examines and manipulates her genitalia briefly and then grasps her hips with his hands and her legs with his feet to mount. Each 5 - 15 second mount involves about ten thrusts. Males sometimes mount two or three times before ejaculating. Infants are whitish-gray, with a distinctive, dark cross pattern along their back and shoulders.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF96FF9CFA0766C7FBA6FB32.taxon	activity	Activity patterns. Banded Langurs are diurnal and arboreal. An activity budget recorded for the Banded Langur was 50 - 2 % resting, 32: 4 % foraging and feeding, 10 - 4 % traveling, 3 - 3 % engaging in social activities, and 3 - 5 % engaging in other miscellaneous activities. Groups usually sleep high up in the forest canopy, sometimes spread out in three to seven differenttrees.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF96FF9CFA0766C7FBA6FB32.taxon	biology_ecology	Movements, Home range and Social organization. Social groups of Banded Langurs are unimale-multifemale and typically of 4 - 10 individuals. Home ranges of four groups averaged 23 - 4 ha (range 16 - 6 - 33 - 1 ha). Daily travel distances averaged 936 m (range 475 - 1500 m).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF96FF9CFA0766C7FBA6FB32.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Near Threatened on The IUCN Red Lust, including the subspecies robinsoni. The nominate subspecies femoralis is classified as Vulnerable and percura as Data Deficient on The IUCN Red List. In 2006, less than 20 individuals were known to remain in Singapore, although juveniles were observed. The overall abundance of the Banded Langur is poorly known. Habitat loss in the form of deforestation and conversion to plantations (mainly oil palm) is the major threat throughout its range. The Banded Langur occurs in at least six protected areas: Raffles’s Banded Langur in Bukit Timah Nature Reserve in Singapore; East Sumatran Banded Langur in Giam Duri Nature Reserve and Siak Kecil Nature Reserve in Sumatra, Indonesia; and Robinson’s Banded Langur in Tanintharyi National Park in Myanmar and Kaeng Krachan National Park and Khao Sok National Park in Thailand. Further studies on the taxonomy, distribution, abundance, and threats to the Banded Langur are urgently needed.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF99FF9CFF9A6007F8CDFC68.taxon	materials_examined	Indonesia, Pontianak, West Borneo.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF99FF9CFF9A6007F8CDFC68.taxon	discussion	An isolated population is found in southeastern Sarawak State (Betong, Saribas, Nimong, Batang Lupar) that probably represents an undescribed subspecies superficially; oo similar to the “ “ cross-bearing 3 of morph of the subspecies cruciger. Two subspecies recognized.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF99FF9CFF9A6007F8CDFC68.taxon	distribution	Subspecies and Distribution. P. c. chrysomelasS. Miller, 1838 — WBorneo, fromtheIVDivisioninWSarawak, StotheKapuasRiverinWKalimantanProvince. P. c. cruciger Thomas, 1892 — N Borneo, from Melalap in Sabah State to the Baram District in N Sarawak.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF99FF9CFF9A6007F8CDFC68.taxon	description	Descriptive notes. Head — body 43 - 61 cm, tail 61 - 84 cm; weight 5.9 - 8.2 kg. Forehead whorls of the Cross-marked Langur, of which there is nearly always a pair, are very small and nearer the brow than in other langur species. Crest is high, upstanding, and narrow, and there is no defined fringe over eyes. There is a U-shaped parting on nape rather than a restricted whorl. Chest hairs are directed backward, meeting forward-directed belly hairs in a ridge. The “ Bornean Banded Langur ” (P. c. chrysomelas) is quite variable but generally jet-black above (sometimes with a few brown hairs on back). Underside is pale whitish or fawn, the tone extending to cheeks, chin, wrists, lower thighs, and ventral surface of tail. Cheeks are brown-gray to white. The “ Tricolored Langur ” (P. c. cruciger) is mainly brown, with a white or creamy-fawn underside (latter tone extending to wrists, ankles, and tail tip). Forehead whorls are relatively elongated, and crest is low. There is also a strikingly different and fairly common “ crossbearing ” morph, in which the characteristic black cross pattern of infants is retained into adulthood. Legs, flanks, and crown are bright red. The isolated population found in south-eastern Sarawak has a less sharp dividing line between the black dorsum and red flanks, a dull creamy underside rather than white, and the chest is a more pronounced red. Black patches on knees are also more conspicuous and there is always black patching on shanks. There are two small whorls on the forehead, with a tall crest between them.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF99FF9CFF9A6007F8CDFC68.taxon	biology_ecology	Habitat. Swamp, lowland, and mangrove forest.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF99FF9CFF9A6007F8CDFC68.taxon	food_feeding	Food and Feeding. The Cross-marked Langur eats leaves and fruit.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF99FF9CFF9A6007F8CDFC68.taxon	breeding	Breeding. Infant Cross-marked Langurs are whitish-gray with a distinctive, dark cross pattern along their back and shoulders.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF99FF9CFF9A6007F8CDFC68.taxon	activity	Activity patterns. The Cross-marked Langur is diurnal and arboreal.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF99FF9CFF9A6007F8CDFC68.taxon	biology_ecology	Movements, Home range and Social organization. The Cross-marked Langur has never been studied in the wild. It generally lives in groups of 3 - 7 individuals, with one male, 1 - 3 females, and their offspring.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF99FF9CFF9A6007F8CDFC68.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Critically Endangered on The IUCN Red Last, including both subspecies. The Cross-marked Languris legally protected in Sarawak. According to the published distributions, the Tricolored Langur occurs in Maludam, Similajau, and Tanjung Datu national parks and Samunsam Wildlife Sanctuary. It is also present in the Lingga area of Sarawak, which has been proposed as a protected area, and it may be found in Betung Kerihun National Park in Indonesia. A recent survey by K. Aken and J. W. Duckworth and colleagues has, however, reported that the Bornean Banded Langur, not the Tricolored Langur, occurs in Similajau National Park, so the respective distributions of the two subspecies (and perhaps their validity as distinct taxa) require further studies. According to its known and supposed distribution, the Bornean Banded Langur may occur in Lanjak-Entimau Wildlife Sanctuary in southwestern Sarawak near the border with West Kalimantan. The majority of the remaining populations are found in these areas, and the combined estimates for their numbers are very low, c. 200 - 500 individuals. Known populations of Cross-marked Langurs are believed to have been reduced by ¢. 80 % over the past 30 years, and certainly the area of occupancy has been reduced dramatically. It was once considered common, but now, after 100 years of intense persecution and habitat loss, it is among the rarest primates in the region, and among the rarest in the world. It occurs in less than 5 % ofits historical distribution. In recent years, it has been particularly affected by expanding plantations, particularly oil palm. Additional conservation measures are urgently needed. The Crossmarked Langur could soon go extinct if current trends continue.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF99FF9CFAE666CDF5B2F3CF.taxon	materials_examined	Indonesia, south-eastern Borneo. Restricted by L. Medway in 1965 to south-eastern Kalimantan, Murung and “ Pulu Lampy, ” near Banjarmasin, Pematang, Kuala.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF99FF9CFAE666CDF5B2F3CF.taxon	discussion	This species is monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF99FF9CFAE666CDF5B2F3CF.taxon	distribution	Distribution. Found patchily in C & E Borneo, from C Sarawak State (absent from coastal areas and possibly occurs only between upper Rajang and upper Batang Luparrivers) to most of E Kalimantan and a few areas in the W (in the SE restricted to the E of the Barito River, there is a population in Gunung Palung in the SW, but it is not known whetherit is isolated or whetherit is continuous with other populations in the W).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF99FF9CFAE666CDF5B2F3CF.taxon	description	Descriptive notes. Head — body 50 - 54 cm (males) and 47 - 50 cm (females), tail 70 - 74 cm (males) and 63 - 70 cm (females); weight ¢. 5 - 7 kg. The White-fronted Langur is pale grayish-brown above, with a yellowish-brown underside. Brow, crown crest, and cheeks are blackish and lower segments of limbs grade toward black of hands and feet. Tail is yellowish-gray. There is a naked patch of white skin on forehead (hence the common name) set off by one or two whorls of hair, with a tall, forward-leaning crown crest behind.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF99FF9CFAE666CDF5B2F3CF.taxon	biology_ecology	Habitat. Tall lowland, riparian and hill dipterocarp forest, up to elevations of 1000 m. The White-fronted Langur may also be found in secondary forest and old rubber tree plantations. Very little is known about its habitat use and ecology.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF99FF9CFAE666CDF5B2F3CF.taxon	food_feeding	Food and Feeding. The White-fronted Langur eats mainly young leaves, supplemented with fruits, seeds, and flowers.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF99FF9CFAE666CDF5B2F3CF.taxon	breeding	Breeding. The natal coat of the White-fronted Langur is white, with a thick black dorsal stripe.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF99FF9CFAE666CDF5B2F3CF.taxon	activity	Activity patterns. The White-fronted Langur is diurnal and arboreal.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF99FF9CFAE666CDF5B2F3CF.taxon	biology_ecology	Movements, Home range and Social organization. The White-fronted Langur has never been studied in the wild. Groups are small, with a single adult male and 2 - 3 females. The home range is estimated to be larger than 100 ha. There are an estimated 13,400 White-fronted Langurs in Sarawak (Lanjak-Entimau), with densities of 1 - 7 groups / km?. Density estimates from ten differentsites are 1 - 5 groups / km? and 5 - 9 ind / km? ®.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF99FF9CFAE666CDF5B2F3CF.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Vulnerable on The IUCN Red List. The White-fronted Langur is protected by Indonesian law and is totally protected in Sarawak. Its distribution is fragmented. It is frequently hunted in Sarawak for its gallstones and meat. Habitat loss for the White-fronted Langur has been very rapid, particularly due to conversion into oil palm plantations. It occurs in at least seven protected areas: Batang Ai National Park and Lanjak-Entimau Wildlife Sanctuary in Sarawak and Betung Kerihun, Bukit Baka-Bukit Raya, Gunung Palung, and Kutai national parks and Sungai Wain Protection Forest in Kalimantan. The level of protection that these areas offer, particularly in Indonesia, is insufficient, and there is a need to reinforce their management. In some areas, such as the north-western mountains (Lanjak-Entimau, Bantang Ai, and Bentuang Kerima), the White-fronted Langur seems to be common, but in other areas, it is evidently scarce.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF99FF9DFAE4696CFB8DF26D.taxon	materials_examined	Malaysia, Sarawak, Niah.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF99FF9DFAE4696CFB8DF26D.taxon	discussion	The forms sabana and canicrus that are commonly considered to be subspecies of P. hosei are in fact distinct species, based primarily on morphological differences. The population at the type locality evidently lacks sexual dichromatism, and on this basis, two subspecies have been distinguished: hosei, supposed to be restricted to the region around Niah (possibly extending east to Brunei), lacking the sexual color difference, and everetti, named by O. Thomas in 1892, from the rest of the distribution of the species (inland eastern Sarawak, Brunei, west Sabah and northern Kalimantan) possessing it. Nevertheless, it is probable that ages of the crucial specimens, on which the distinction was made, were misunderstood; that is, they were young females instead of adults. The facial skin of P. hosei is pinkish, except for black around eyes and bluish lips. Considered monotypic here.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF99FF9DFAE4696CFB8DF26D.taxon	distribution	Distribution. NW Borneo, W Sabah State (along W coast N to Mt Kinabalu), NE Brunei, and N Sarawak State (from its type locality Niah and the lower Baram River, E inland to 2 ° 40 ’ N, and the border with N Kalimantan).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF99FF9DFAE4696CFB8DF26D.taxon	description	Descriptive notes. Head-body 48 - 56 cm, tail 65 - 84 cm; weight 6 - 7 kg (males) and 5.5 - 6 kg (females). Hose’s Languris iron-gray, with whitish tips on its upper side, giving an overall grizzled effect. Underside is grayish to buffy-white, becoming white on inner sides of limbs, and hands and feet are black — this tone sometimes extending to lower parts of legs. Crown crest is black, with the remainder of the crown sometimes being blackish. Facial skin is pinkish on lower jaw and cheeks, and darker (reddish or black) elsewhere. Upperparts are uniformly grayish-black. Legs are darker distally, becoming black on feet. Tail is paler below than above. Hose’s Langur is sexually dichromatic, the male having the black color restricted to its crown, with the rest of the head being white, whereas in the female, the black area also covers nape and forehead, extending laterally to ears. Juveniles are colored like adult males; typical pattern of the adult female develops at maturity.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF99FF9DFAE4696CFB8DF26D.taxon	biology_ecology	Habitat. Lowland to hill dipterocarp rainforest (Mount Kinabalu). Elevational range is sea level to ¢. 1000 m, with occasional records to 1600 m (e. g. Crocker Range) and possibly higher. Hose’s Langurs occasionally enter plantations. They are sympatric with the Maroon Langur (P. rubicunda) in the northern part of their distribution, the Crossmarked Langur (P. chrysomelas) in the western part, and the White-fronted Langur (P. frontata) elsewhere; but one or the other at any locality is absent or rare.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF99FF9DFAE4696CFB8DF26D.taxon	food_feeding	Food and Feeding. Hose’s Langurs eat mainly young leaves, along with unripe fruits, seeds, flowers, bird eggs, and nestlings. Individuals churn up and eat mud around mineral springs, which results in the formation of stomach concretions known as “ bezoars ” that are sought after by local people for their supposed medicinal properties.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF99FF9DFAE4696CFB8DF26D.taxon	breeding	Breeding. Infant Hose’s Langurs are white, with a black cruciform pattern across their backs and shoulders.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF99FF9DFAE4696CFB8DF26D.taxon	activity	Activity patterns. Hose’s Languris diurnal and arboreal.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF99FF9DFAE4696CFB8DF26D.taxon	biology_ecology	Movements, Home range and Social organization. Hose’s Languris territorial and lives in groups with one male, 2 - 6 females, and their young. In Brunei, the average group size 1 s six. Males produce loud-calls, spacing vocalizations, which carry some distance; they are simpler in structure than those of other species of Presbytus.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF99FF9DFAE4696CFB8DF26D.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Data Deficient on The IUCN Red List (as P. hosei hose). Although no subspecies are recognized here, the IUCN assessment included the form everetti as Vulnerable. Hose’s Langur is legally protected in Indonesia, although wrongly named P. aygula, a misidentification that may serve to subvert prosecution of offenders. P. aygula is the former name for the Javan Langur (P. comata), a species with which Hose’s Langur has sometimes been incorrectly lumped in the past; the original description of Simia aygula, however, was based on the long-tailed or crab-eating macaque, not on a langur at all. The conservation status of Hose’s Langur needs to be reassessed. If the sexually dichromatic form everetti is valid, then it is believed that the form hose: (sexually monochromatic) would be restricted to the northern part of the Similajau National Park (a 90 km strip of forest along the Sarawak coast and now completely isolated by oil palm plantations) and possibly Brunei a little to the north-east. There is, to date, no evidence of its occurrence in Similajau (although the possibility still remains; white-whiskered individuals have been seen, but their identification is inconclusive) and surveys in Brunei are needed to confirm its continued existence there. Hose’s Languris relatively tolerant of forest disturbance and light logging, but the destruction of its forests and hunting are ongoing threats. In 2010, it was estimated that their forests had been reduced by more than 50 % by settlements and primarily monoculture plantations for wood pulp and palm oil. Widespread and intense forest fires have also been problematic; they result from the drying of the forest floor because of selective (intensive) logging are serious during El Nino years. About 50,000 km? of its forests remain. The hunting of Hose’s Langurs correlates with the presence of inland settlements, the timber industry, and the demand for lucrative bezoars (a visceral mass believed to be a poison antidote that is traded to supply the traditional Chinese medicine market). Hose’s Langurs are also hunted for food and to deter crop raiding. Most of the presumed (possible) distribution of Hose’s Langur coincides with oilfields that straddle the frontier between Sarawak and Brunei. Hose’s Langur (including everett) occurs in Mount Kinabalu National Park and Lanjak-Entimau Wildlife Sanctuary in Sarawak and Sungai Kayan National Park and Betung-Kerihun National Park in Indonesia. There are no Hose’s Langurs in captivity.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF98FF9DFA036405F7DCF5F4.taxon	materials_examined	Indonesia, East Kalimantan (Dutch north-east Borneo).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF98FF9DFA036405F7DCF5F4.taxon	discussion	This species is monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF98FF9DFA036405F7DCF5F4.taxon	distribution	Distribution. Borneo, confined to an area of E Kalimantan, from the Kayan and Sembakung (= Sebuku) rivers in the N, S, and SW to the Mahakam River, the W boundary is not well known but certainly E of the Apau Kayan and the Kayan Mentarang National Park.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF98FF9DFA036405F7DCF5F4.taxon	description	Descriptive notes. eds 48 - 56 cm, tail 65 - 84 cm; weight 6 - 7 kg (males) and 5.5 - 6 kg (females). Hairs of upperparts of Miller’s Langur are iron-gray with whitish tips, giving an overall grizzled effect. Underside is grayish to buffy-white, turning white on inner sides of limbs, and hands and feet are black — this tone sometimes extending to lower parts of legs. Crown crest is black, with remainder of crown sometimes blackish. Facial skin is pinkish on lower jaw and cheeks, and darker (reddish or black) elsewhere. Miller’s Langur is a small, long-tailed form, generally similar to the Sabah Grizzled Langur (P. sabana), but crown is wholly brownish and underside is buffy-white; its facial skin is blackish down to a line between mouth and ears, at which point it becomes whitish-buff.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF98FF9DFA036405F7DCF5F4.taxon	biology_ecology	Habitat. Lowland to hill dipterocarp rainforest. Elevational range is sea level up to c. 1000 m, with occasional records up to 1600 m and possibly higher. Miller’s Langur (referred to at the time as P. aygula) was studied by P. Rodman in lowland (up to 300 m) dipterocarp forest and riparian forest along the Sengata River in Kutai National Park in East Kalimantan. Densities were slightly lower in riparian forest (18 - 4 ind / km?) compared to interior dryland forest (22 - 3 ind / km?). It was found to be sympatric with two other langurs, the White-fronted Langur (PF. frontata) and the Maroon Langur (P. rubicunda), which were, however, scarce compared with Miller’s Langurs.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF98FF9DFA036405F7DCF5F4.taxon	food_feeding	Food and Feeding. Miller's Langurs eat mainly young leaves and leaf shoots (66 %), along with unripe fruits (28 %); also seeds, flowers, bird eggs, and nestlings. Individuals churn up and eat mud around mineral springs, resulting in the formation of stomach concretions known as “ bezoars, ” which are sought after for their supposed medicinal properties.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF98FF9DFA036405F7DCF5F4.taxon	breeding	Breeding. Infant Miller’s Langurs are white, with a black cruciform pattern across their backs and shoulders.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF98FF9DFA036405F7DCF5F4.taxon	activity	Activity patterns. Miller's Langur is diurnal and arboreal. The majority of its time is spent in the middle and upper forest canopies above heights of 20 m (62 %), and 29 % of its time is spent in the lower canopy, 10 - 20 m above the ground.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF98FF9DFA036405F7DCF5F4.taxon	biology_ecology	Movements, Home range and Social organization. Social groups of Miller’s Langurs have a single male, 2 - 4 females, and their young. Mean group size is eight individuals.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF98FF9DFA036405F7DCF5F4.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Endangered on The IUCN Red List (as P. hosei canicrus). Miller’s Langur is legally protected in Indonesia, under the name P. aygula canicrus. The geographical boundaries ofits distribution are unknown. The formerly large population in Kutai National Park is considerably reduced due to widespread and repeated forest fires during El Nino years in the 1980 s and 1990 s. A previously unknown population was discovered in Wehea Forest, further inland, in the 2000 s. In this forest, camera-trap records at one salt lick were only 0 - 2 Miller’s Langurs / day, similar to Bornean Orangutans (Pongo pygmaeus), and 50 % that of Maroon Langurs. At a different salt lick, there were 0 - 7 Miller’s Langurs / day, and they were often the most recorded primate. This forest, fortunately, is protected by provincial and local (customary) law. There are no Miller’s Langurs in captivity.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF98FF9EFA036F44FB54F7A6.taxon	materials_examined	Malaysia, Sabah, Paitan.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF98FF9EFA036F44FB54F7A6.taxon	discussion	Until recently, P. sabana was classified as a subspecies of P. hoser, but it is morphologically distinct. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF98FF9EFA036F44FB54F7A6.taxon	distribution	Distribution. N Borneo in most of Sabah State, as far SW as Kalabakan (4 ° 26 ’ N 117 ° 29 ’ E); its occurrence in N Kalimantan is not confirmed.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF98FF9EFA036F44FB54F7A6.taxon	description	Descriptive notes. Head-body 48 - 56 cm, tail 65 - 84 cm; weight 6 - 7 kg (males) and 5. 5 - 6 kg (females). The Sabah Grizzled Languris gray above and below, except white belly and inner surfaces of limbs, contrasting with black on hands and feet. Crown is gray, with a tall black crest on back of head and two frontal whorls. Facial skin is reddish, with bluish lips and a pair of distinct black spots on each side of the face, between eyes and mouth — a pattern sometimes seen in more contrastingly colored individuals of Hose’s Langur (Presbytis hoset).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF98FF9EFA036F44FB54F7A6.taxon	biology_ecology	Habitat. Evergreen, lowland, riparian, and montane rainforest. Sabah Grizzled Langurs will also enter tree plantations.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF98FF9EFA036F44FB54F7A6.taxon	food_feeding	Food and Feeding. Sabah Grizzled Langurs eat mainly young leaves, unripe fruits, seeds, and flowers. Diets vary with the extent of habitat degradation (logging), in different forests, and at different times of the year. On average, young leaves comprise 42 - 45 % ofthe diet, fruits 3 - 25 %, seeds 17 - 21 %, mature leaves only 1 - 5 %, and flowers 3 % or less. They also eat buds, bark, insects, and bird eggs, and nestlings. More leaves, especially those oflianas (mostly Fabaceae), are eaten in unlogged forest (78 % of the diet vs. 60 % in logged forest), and more fruits and seeds are eaten in logged forest (19 % fruits and 21 % seeds vs. 3 % fruits and 17 % seeds in unlogged forest).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF98FF9EFA036F44FB54F7A6.taxon	breeding	Breeding. Infants are weaned at ¢. 300 days; they are consideredjuvenile at c. 11 months of age and subadult at 21 - 36 months. Infants are white, with a black cruciform pattern across their backs and shoulders. Females first give birth at ¢. 3 years old and usually in July-October.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF98FF9EFA036F44FB54F7A6.taxon	activity	Activity patterns. The Sabah Grizzled Langur is diurnal and arboreal. As for other species of Presbytis, resting dominates the activity budget, necessary to ferment and digest their leafy diet. Groups feed and forage for c. 14 - 17 % of the day, rest for 45 - 49 %, travel for 30 - 32 %, and indulge in social activities for 4 - 7 - 6 %. They travel and forage mostly in the middle and upper forest canopy at heights of 10 - 30 m. They sleep high in the middle and upper canopies of emergent trees. Their locomotion involves quadrupedal walking (¢. 47 %), climbing (c. 22 %), running (c. 12 %), occasional leaping (c. 10 %), vertical clinging and leaping (c. 7 %), and rarely suspensory swinging (c. 1 %) and hopping (c. 1 %).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF98FF9EFA036F44FB54F7A6.taxon	biology_ecology	Movements, Home range and Social organization. The Sabah Grizzled Langur is territorial and lives in one-male groups of up to 14 individuals, with 2 - 6 females and their young. Home ranges vary from ¢. 35 ha to 125 ha, and daily travel distances are ¢. 700 - 800 m. Males produce loud calls that are spacing vocalizations and carry some distance. They are simpler in structure than is typical of Presbytis and are similar to those of Hose’s Langur, differing only in a few details. Long calls or “ spacing calls, ” as they are called, are described as “ gargling growls. ” When groups meet, males become agitated; they run and leap, uttering series of these gargling growls. Their alarm calls for predators are staccato grunts. Females disperse from their natal groups. Densities are estimated at 2 - 1 groups / km?, or ¢. 16 ind / km? ®. They occasionally form mixed species groups with Maroon Langurs (P. rubicunda), although where one is present, the other tends to be scarce or absent.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF98FF9EFA036F44FB54F7A6.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Endangered on The IUCN Red List (as P. hosei sabana). The Sabah Grizzled Languris protected by Indonesian law under the name P. aygula. Hunting for food and bezoars, believed to have medicinal properties, is its major threat, and believed to be the principal cause for its decline. The Sabah Grizzled Langur seems to survive selective, light logging quite well, but logging that destroys the forest canopy (and eliminates their sleeping trees) and clearcutting are major threats. It occurs in Danum Valley Conservation Area, Tabin Reserve, Sepilok Orang Utan Sanctuary, and Lower Kinabatangan and Maliau Basin conservation areas in Sabah. It may also occur in Crocker Range Park. There are no reports of the Sabah Grizzled Langur in captivity.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9BFF9EFF916DB3F9D9F8B8.taxon	materials_examined	Indonesia, Bencoolen (= Bengkulu), Sumatra.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9BFF9EFF916DB3F9D9F8B8.taxon	discussion	Formerly, P. mitrata, P. bicolor, and P. sumatrana were considered subspecies of P. melalophos. There are widespread geographic gradients in color and individual variation (often in the form of wellmarked morphs), making assignment of subspecies difficult. In addition to the typical P. melalophos, there are three main morphs that coexist widely overits distribution. They have been described as “ nobilis ” (foxy-red), “ ferruginea’ (less reddish), and “ aurata’ (golden-buff with a black spot on chest, black hands and feet, and a white tail-tip). There is a fourth form described by N. F. Chasen in 1940 with the name fluviatilis. It is found in Palembang, Sumatra, and is similar to the Black-crested Sumatran Langur but much paler. The taxonomic status of these morphsis currently under study. According to genetic and acoustic data, the proposed distribution of P. melalophos may have to be extended into the distribution of P. mitrata. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9BFF9EFF916DB3F9D9F8B8.taxon	distribution	Distribution. W Sumatra, from the upper Rokan River (Tanangtalu, 0 ° 20 " N, 99 ° 59 ’ E), just N of Gunung Talakmau, S to the upper Batang Hari River and beyond along the Barisan Range, W of Lampung Province.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9BFF9EFF916DB3F9D9F8B8.taxon	description	Descriptive notes. Head — body 34: 1 - 39.7 cm (males, n = 15) and 25.7 - 40.4 cm (females, n = 13), tail 68 - 80 cm (males, n = 15) and 67.5 - 86 cm (females, n = 13); weight 5.9 - 9 kg (males, n = 15) and 5.2 - 8.9 kg (females, n = 14). The Black-crested Sumatran Languris variable in its pelage and coloration. It is so-named for the distinctive, blackpointed crest on its crown. Coat color ranges from white through gray to reddishorange. Tail is normally bi-colored (dark above and white below). The Black-crested Sumatran Langur is generally reddish-orange above (tipped with black), with a white, yellow, or pale orange underside. Outersides of limbs tend to be paler (more orange), and inner sides of limbs white, with hands and feet usually orange but sometimes black. Tail is like the body above and whitish underneath for the proximal half. Facial skin is black, and cheeks are straw-colored and separated from the whitish forehead by a reddish or blackish band. Crown crest is black, and there is usually a single, indistinct whorl on forehead (none or two in some populations), as well as a U-shaped parting on nape.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9BFF9EFF916DB3F9D9F8B8.taxon	biology_ecology	Habitat. Primary and secondary lowland and hilly rainforest, rubber plantations, and bush areas. The Black-crested Sumatran Langur uses all levels of the forest canopy. It tolerates a certain degree of habitat degradation and can be found in disturbed areas and secondary forests.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9BFF9EFF916DB3F9D9F8B8.taxon	food_feeding	Food and Feeding. Black-crested Sumatran Langurs eat young leaves, unripe fruits, flowers (e. g. Erythrina indica, Fabaceae), mature leaves, and seeds. In Bukit Sebelah Protection Forest, much of the diet consists of leaves of Hevea brasiliensis (Euphorbiaceae) and Ficus variegata (Moraceae), and they feed mainly at 15 - 20 m above the ground.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9BFF9EFF916DB3F9D9F8B8.taxon	breeding	Breeding. Little information is available for the Black-crested Sumatran Langurs. Allomothering is rare. Newborns are whitish-gray, with a thick, dark dorsal stripe, but no shoulder cross-stripe.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9BFF9EFF916DB3F9D9F8B8.taxon	activity	Activity patterns. The Black-crested Sumatran Langur is diurnal and arboreal.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9BFF9EFF916DB3F9D9F8B8.taxon	biology_ecology	Movements, Home range and Social organization. Home ranges of Black-crested Sumatran Langurs are 14 - 30 ha, and daily movements are ¢. 300 - 1360 m. Their social groups are small, with 2 - 8 individuals and a single adult male. The Black-crested Sumatran Langur has never been studied in the wild.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9BFF9EFF916DB3F9D9F8B8.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Near Threatened on The IUCN Red List (as P. melalophos melalophos). The Black-crested Sumatran Langur is protected by Indonesian law. There has been widespread recent loss of habitat in its distribution, mainly for oil palm plantations. Nevertheless, it is still relatively common, and it seems thatit can tolerate some degree of land conversion and forest degradation. Populations are now, however, very fragmented. It is commonly trapped for the illegal pet trade, but less so than the Mitered Langur (P. mitrata). This and some incidental hunting for food are threats acrossits range. It occurs in Kerinci-Seblat National Park and Bukit Sebelah Protection Forest in Sumatra. The still rather confused taxonomy, confounded by different color variants and little understood boundaries of the ranges of this and other species (formerly considered subspecies), requires further research to better understand the conservation status of the Black-crested Sumatran Langur.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9BFF9FFA9A629DFC13FE72.taxon	materials_examined	Batang Kering 0 ° 50 ° S, 101 ° 23 ° E, 14 km north of Kiliranjao, West Sumatra.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9BFF9FFA9A629DFC13FE72.taxon	discussion	P. bicolor is aligned with P. melalophos on the basis of similarity of vocalizations and genetics but with P. femoralis or P. siamensis based on pelage color. It is separated from P. melalophos by the middle and lower Batang Hari River (although intergrading with it in foothills to the west), and from P. siamensis by the middle Inderagiri River. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9BFF9FFA9A629DFC13FE72.taxon	distribution	Distribution. EC Sumatra, from the Inderagiri River in the N to the Batang Hari River in the S, mainly in the highlands.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9BFF9FFA9A629DFC13FE72.taxon	description	Descriptive notes. Head-body 33 - 35.4 cm (males) and 33.9 - 38.2 cm (females), tail 64 - 72.5 cm (males) and 66 - 84 cm (females); weight 4.9 - 7.2 kg (males) and 5 - 1 - 8: 5 kg (females). Main body color of the Black-and-white Langur is dark chocolate, extending down outer surfaces of arms. Upperpart of tail and outer surfaces of hindlimbs, hands, and feet are black. Throat, underside, inner surfaces of limbs, and undersurfaces oftail are white. Crest on crown has a black median stripe and a black tip. There is a black fringe of hair over forehead. Circumocular region is gray or bluishgray, muzzle is black, and chin is pinkish or gray.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9BFF9FFA9A629DFC13FE72.taxon	biology_ecology	Habitat. Mainly hill rainforest.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9BFF9FFA9A629DFC13FE72.taxon	food_feeding	Food and Feeding. The diet of the Black-and-white Langur probably consists of young leaves, unripe fruits, flowers, mature leaves, and seeds.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9BFF9FFA9A629DFC13FE72.taxon	breeding	Breeding. Newborn Black-and-white Langurs are whitish-gray with a thick, dark dorsal stripe, and no shoulder cross-stripe.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9BFF9FFA9A629DFC13FE72.taxon	activity	Activity patterns. The Black-and-white Langur is diurnal and arboreal.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9BFF9FFA9A629DFC13FE72.taxon	biology_ecology	Movements, Home range and Social organization. Like other species of Presbytis, the Black-and-white Langur likely lives in one-male groups and defends exclusive territories.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9BFF9FFA9A629DFC13FE72.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Data Deficient on The [UCN Red List (as P. melolophos bicolor). The Black-and-white Langur is protected under Indonesian law. It occurs in Bukit Tigapuluh National Park. Further surveys and clarification are needed to assessits distribution and conservation status.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9AFF9FFA2D6405F9D9F783.taxon	materials_examined	Indonesia, Sumatra, mainland opposite Zutphen Islands.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9AFF9FFA2D6405F9D9F783.taxon	discussion	P. mitrata was formerly included in P. melalophos as subspecies. There are widespread geographic gradients in color and individual variation. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9AFF9FFA2D6405F9D9F783.taxon	distribution	Distribution. SE Sumatra E of the Barisan Range, from the Batang Hari River, SW to the upper Musi River drainage, and S to Lampung Province.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9AFF9FFA2D6405F9D9F783.taxon	description	Descriptive notes. eniionds 42 - 57 cm (males) and 42 - 56 cm (females), tail 64 — 82 cm (males) and 62 - 82 cm (females); weight ¢. 5 - 9 kg (males) and c. 5 - 8 kg (females). Upper body of the Mitered Langur (including forehead and crown) ranges from dark mouse-brown through ashy-gray to pale red-yellow or yellow-gray, with a blackish overlay. Underside is creamy-yellow or white (extending well up onto flanks), limbs are mixed whitish and gray or red-brown, hands and feet are gray, and tail is redder than body above and pale buffy below, with a white tip. There are generally no whorls, but a black D-pattern on crest encloses white hairs of fore-crown, and thereis a white cheek ruff. Face is gray, and muzzle is often pink, with a white or pink crescent present around outer corners of eyes. There is a mostly white morph that has been ascribed to this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9AFF9FFA2D6405F9D9F783.taxon	biology_ecology	Habitat. Primary and secondary lowland rainforest, rubber plantations, and bushy areas. Mitered Langurs will use all levels of the forest canopy but tends to favor the understory. It is tolerant of habitat conversion to a degree and can be found in disturbed and secondary forest areas.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9AFF9FFA2D6405F9D9F783.taxon	food_feeding	Food and Feeding. Mitered Langurs eat young leaves, unripe fruits, flowers, mature leaves, and seeds.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9AFF9FFA2D6405F9D9F783.taxon	breeding	Breeding. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9AFF9FFA2D6405F9D9F783.taxon	activity	Activity patterns. The Mitered Languris diurnal and arboreal.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9AFF9FFA2D6405F9D9F783.taxon	biology_ecology	Movements, Home range and Social organization. Home ranges of Mitered Langurs are 14 - 29 - 5 ha. The daily distance moved is c. 300 - 1360 m.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9AFF9FFA2D6405F9D9F783.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Endangered on The IUCN Red List (as P. melalophos mitrata). The Mitered Languris protected by Indonesian law. Habitat loss, especially due to oil palm plantations, is a major threat, but they have some tolerance to forest conversion. Like the Black-crested Sumatran Langur (P. melalophos), the Mitered Langur is relatively common in remaining habitat, but its occurrence is very patchy and fragmented. It is heavily hunted and often trapped for the pet trade. It occurs in Bukit Barisan Selatan National Park and Way Kambas National Park in Sumatra. Further study is urgently needed to better understand its distribution and the geographic patterns of color variants.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9AFF9FFF2164CFFEC4F787.taxon	materials_examined	“ Mt. Ophir ” (= G. Talamau), north of Padang.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9AFF9FFF2164CFFEC4F787.taxon	discussion	P. sumatrana was formerly considered a subspecies of P. melalophos. Northern populations are darker and blacker than those in the south, which are more brownish, and could be separated as the subspecies margae. It is separated from P. femoralis percura by the Rokan River and from P. siamensis paenulata by the Barumun River. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9AFF9FFF2164CFFEC4F787.taxon	distribution	Distribution. W & NC Sumatra (N highlands and W coast S of the Simpang Kiri River, N of Gunung Talamau, and on the E coast from the Wampu River to the Barumun River, and SE to the Rokan River), and Batu Is (Pini I).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9AFF9FFF2164CFFEC4F787.taxon	description	Descriptive notes. Head — body 33 - 35 cm (males) and 34 - 39 cm (females), tail 64 — 72 cm (males) and 66 - 84 cm (females); weight 4.9 - 7.2 kg (males) and 5.1 - 8.5 kg (females). Upperparts of the Black Sumatran Langur are dark gray-brown to nearly black, including dorsal surface of tail. Hands, feet, and outer limbs are black; throat, underparts, underside oftail, and inner surfaces of limbs as far as wrists and ankles are white, sharply set off from dark color of upperparts. Crest is gray to dark brown, and sometimes indistinctly formed. Facial skin is mostly bluish, flesh colored around the mouth, but lips are black.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9AFF9FFF2164CFFEC4F787.taxon	biology_ecology	Habitat. [Lowland and hill rainforest, old growth forest, and secondary forest.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9AFF9FFF2164CFFEC4F787.taxon	food_feeding	Food and Feeding. The diet of the Black Sumatran Langur probably consists largely of young leaves, unripe fruits, flowers, mature leaves, and seeds.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9AFF9FFF2164CFFEC4F787.taxon	breeding	Breeding. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9AFF9FFF2164CFFEC4F787.taxon	activity	Activity patterns. The Black Sumatran Langur is diurnal and arboreal.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9AFF9FFF2164CFFEC4F787.taxon	biology_ecology	Movements, Home range and Social organization. The Black Sumatran Langur has never been studied in the wild. Like other species of Presbytis, it likely lives in small onemale groups and defends exclusive territories.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9AFF9FFF2164CFFEC4F787.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Endangered on The IUCN Red List (as P. melalophos sumatranus). The Black Sumatran Langur is protected under Indonesian law. About 50 % of the forests in its range have been degraded over the past 30 years. Remaining forest is a fragmented mosaic among commercial plantations. The illegal pet trade is a further threat to the Black Sumatran Langur. It occurs in Batang Gadis National Park.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9CFF99FF2D6401F6EDFC46.taxon	materials_examined	Tenasserim. Corrected to Mentawai Islands by O. Thomas in 1895, and restricted to Sipora Island by F. N. Chasen and C. B. Kloss in 1927. Amended by D. Brandon-Jones in 1993 to Pulau Pagai Utara (= North Pagai Island).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9CFF99FF2D6401F6EDFC46.taxon	discussion	Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9CFF99FF2D6401F6EDFC46.taxon	distribution	Distribution. Mentawai Is (Sipora, North Pagai, and South Pagai Is, also on the islet of Sinakak off the E coast of South Pagai).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9CFF99FF2D6401F6EDFC46.taxon	description	Descriptive notes. Head-body c. 50 cm, tail c. 58 cm; weight c. 6 - 5 kg (males) and c. 6 - 4 kg (females). The Mentawai Langur is jet-black above and on tail, with underside and innersides of thighs reddish-orange. Innersides of upper arms are whitish or reddish, and throat, cheeks, and chin are yellowish-white, grading backward through gray to the reddish color of underside. Brow band is whitish. Pubic region is yellowish-white in both sexes, and males have a white scrotum. Facial skin is black, becoming somewhat depigmented around the mouth. There is a small, erect, slightly forward-directed crest on crown and a ridge around chin and up to ears, where the forward-directed interramal hair stream meets the backward-directed cheek hair stream. Underparts are bright reddish-orange, with black-tipped hairs toward the midline of the chest. There may be slight differences between individuals on Sipora and the Pagaiislands.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9CFF99FF2D6401F6EDFC46.taxon	biology_ecology	Habitat. Primary and secondary lowland evergreen rainforest, swamps, logged forest, and (possibly) mangrove forest and cultivated areas. Mentawai forests are ever-wet rainforests, and annual rainfall is up to 4000 mm. Mentawai Langurs use all levels of the forest canopy, even occasionally coming to the ground, but they spend most of their active time in the middle canopy and resting time in the upper canopy. They are most abundant in forest logged 20 years ago and least abundant in recently logged forest.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9CFF99FF2D6401F6EDFC46.taxon	food_feeding	Food and Feeding. The Mentawai Langur is highly frugivorous for a colobine, although the proportion of the diet that is fruit and seeds appears to vary with habitat. In one study in a primary forest habitat on North Pagai, fruit made up only 32 % of the diet, while in a second study in secondary forest habitat, also on North Pagai, fruit and seeds contributed 70 % of the diet, more than for any other species of colobine observed. Leaves can still make up a significant portion of the diet (22 - 55 % of diet). The Mentawai Langur also eats flowers, sap, bark, and fungi, and it is one of the few primates that can eat leaves of the dipterocarp tree family.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9CFF99FF2D6401F6EDFC46.taxon	breeding	Breeding. Mating of the Mentawai Langur is ventral-dorsal and solicited by the female about two-thirds of the time. A single infant is born every other year in July-August. Infants have white fur and pale skin and attain adultlike coat and skin color by 12 - 14 weeks of age. Infant “ parking ” (mother leaving the infant alone in a tree while she forages) has been observed.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9CFF99FF2D6401F6EDFC46.taxon	activity	Activity patterns. The Mentawai Languris diurnal and arboreal. Morning loud calls by adult males begin at 03: 00 - 05: 00 h. They leave sleeping trees before dawn, earlier than sympatric Kloss’s Gibbons (Hylobates klossii), perhaps to avoid feeding competition. In one study on North Pagai, observations of group movements in early mornings and late evenings suggest that Mentawai Langurs use sleeping trees in coconut groves and swamp forest outside the primary forest, and travel to the primary forest area to feed in the daytime. This movement pattern may be a way to avoid competition for sleeping trees with Kloss’s Gibbons. Group members usually stay within 30 m of each other while feeding and 10 m while resting. About 25 - 33 % of their time is spent feeding and 45 - 56 % resting. Only a very low percentage of time is spentin social activities (1 - 2 %), and grooming is rarely observed.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9CFF99FF2D6401F6EDFC46.taxon	biology_ecology	Movements, Home range and Social organization. On North Pagai, home ranges of the Mentawai Langur are 17 - 40 ha. Social organization is variable; Mentawai Langurs have been observed in unimale — unifemale, unimale — multifemale groups, and multimale-multifemale groups. In areas with high densities and smaller home ranges, males defend territories from other groups, using visual displays and loud vocalizations, sometimes accompanied by adult females in duets. In areas with larger home ranges, adjacent territories may overlap, and groups tolerate each other. Adult males and sometimes females also give loud vocalizations to warn others of predators (people and pythons). Larger group sizes of Mentawai Langurs (compared with the closely related Siberut Langurs, P. siberu) may be an adaptation to increased hunting and the need for increased vigilance. Adult males and females seem to share equal responsibility for leading group movements. Intragroup aggression is infrequent. Densities are c. 1 - 7 groups / km?. It is sympatric with the Pagai Macaque (Macaca pagensis), the Pigtailed Langur (Simias concolor), and Kloss’s Gibbon.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9CFF99FF2D6401F6EDFC46.taxon	conservation	Status and Conservation. CITES Appendix I. Classified as Critically Endangered on The IUCN Red List (as P. potenziani potenziani). The Mentawai Langur is protected by Indonesian law. It is threatened by logging and hunting and is reported to be the second-most popular game species after the Pig-tailed Langur. On Siberut, where its sister taxon, the Siberut Langur, is found, local people are reported to climb up into their sleeping trees at night to shoot them; this hunting method may also be used for Mentawai Langurs. In recent years, hunting has intensified because of increased access to remote areas due to logging roads and tracks and the replacement of bows and arrows with rifles. Local rituals and taboos that formerly regulated hunting have been lost because of the Christian religion. Mentawai Langurs are occasionally kept as pets but do not thrive in captivity. Other threats include oil palm plantations, forest clearing, and product extraction by local people. Population decline of the Mentawai Langur is estimated at more than 50 % over the past 40 years. In 2005, the population was thought to number only 300 - 1200 individuals across the three islands. It is not found in any protected areas, but was present in the proposed Betumonga Research Area, which was logged by 2003. In 2006, the following conservation actions were suggested: protection of areas in the Pagai Islands by cooperating with a logging corporation that has practiced sustainable logging technique there since 1971; conservation education, especially regarding hunting; and the development of alternative economic models for the local people to reduce the likelihood of them selling off their lands to logging companies.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9CFF9AFA3466EEFC94FC6B.taxon	materials_examined	Siberut Island, Indonesia.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9CFF9AFA3466EEFC94FC6B.taxon	discussion	P. siberu is a separate species from P. potenziani on the basis of genetic divergence. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9CFF9AFA3466EEFC94FC6B.taxon	distribution	Distribution. Mentawai Is (Siberut I).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9CFF9AFA3466EEFC94FC6B.taxon	description	Descriptive notes. Head-body 48 - 50 cm, tail 58 - 64 cm; weight c. 6 - 5 kg (males) and c. 6 - 4 kg (females). Thereislittle sexual dimorphism in the Siberut Langur. Ischial callosities are connected in males but separated in females. Adults are predominantly black, with whitish tufts of hair around their faces, atop their heads, around their necks, and on their pubic region in both sexes. Chest, abdomen, and inner sides of thighs are reddish-brown or reddish-orange. Males have a white scrotum. Facialskin is black, becoming somewhat lighter around the mouth. There is a small, erect, slightly forward-directed crest on crown. Coloration is similar to the Mentawai Langur (Presbytis potenziani), but underparts of the Siberut Langur are darker and white pubic patch is more sharply demarcated.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9CFF9AFA3466EEFC94FC6B.taxon	biology_ecology	Habitat. Primary and secondary lowland evergreen rainforest, swamps, logged forests, and (possibly) mangrove forest and cultivated areas. Mentawai forests are ever-wet rainforests, and annual rainfall is up to 4000 mm. Most oftheir active time is spent in the middle canopy and they rest in the upper canopy.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9CFF9AFA3466EEFC94FC6B.taxon	food_feeding	Food and Feeding. The diet of the Siberut Langur has not been studied, but it is probably similar to that of the Mentawai Langur, which is highly frugivorous for a colobine, but also highly folivorous.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9CFF9AFA3466EEFC94FC6B.taxon	breeding	Breeding. Infants have white fur and pale skin but attain adultlike coat and skin color by 12 - 14 weeks of age.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9CFF9AFA3466EEFC94FC6B.taxon	activity	Activity patterns. The Siberut Langur is diurnal and arboreal. Morning loud calls by adult males begin at 03: 00 - 05: 00 h. They leave their sleeping trees before dawn, earlier than sympatric Kloss’s Gibbons (Hylobates klossii), perhaps to avoid feeding competition. Group members usually stay within 30 m of each other while feeding and 10 m while resting. About 25 - 33 % oftheir time is spent feeding and 45 - 56 % resting. Only a very low proportion of time is spent in social activities (1 - 2 %), and grooming is rarely observed.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9CFF9AFA3466EEFC94FC6B.taxon	biology_ecology	Movements, Home range and Social organization. Home ranges of Siberut Langurs are 11 - 5 - 22 ha, somewhat smaller than is typical of the Mentawai Langur. Only unimale — unifemale groups have been observed, unlike the variable social organization observed in the Mentawai Langur. In areas with high population densities and smaller home ranges, males may defend territories from other groups, using visual displays and loud vocalizations, sometimes accompanied by adult females in duets. Adult males and sometimes females also give loud vocalizations to warn others of predators (people and pythons). Adult males and females seem to share equal responsibility for leading group movements. Intragroup aggression is infrequent. A population density of 8 - 2 ind / km? ® was reported at a site in northern Siberut. The Siberut Langur is sympatric with the Siberut Macaque (Macaca siberu), the Pig-tailed Langur (Simias concolor), and Kloss’s Gibbon.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9CFF9AFA3466EEFC94FC6B.taxon	conservation	Status and Conservation. CITES Appendix I. Classified as Endangered on The IUCN Red Last (as P. potenziani siberu). The Siberut Langur is protected by Indonesian law, but its new species designation may not yet be recognized. It is threatened by logging and hunting and is reported to be the second-most popular game animal among Mentawai huntersafter the Pig-tailed Langur. Local people climb up into sleeping trees at night to shoot them. In recent years, hunting pressure has intensified because of increased access to remote areas due to logging roads and tracks and the replacement of bows and arrows with rifles. Local rituals and taboos that formerly regulated hunting have been lost because of the Christian religion. They are occasionally kept as pets but do not thrive in captivity. Other threats include oil palm plantations, forest clearing, and product extraction by local people. Population decline is estimated at more than 50 % over the past 40 years. In 2006, the population was estimated at 1600 - 9500 individuals. The Siberut Langur occurs in only one protected area, Siberut National Park. It is also found in Peleonan Forest in northern Siberut, which, as of 2011, is protected on a short-term basis by local agreements to exclude the area from logging concessions. In 2006, the following conservation actions were suggested: increased protection for Siberut National Park, which currently lacks enforcement; formal protection of Peleonan Forest in northern Siberut, which has unusually large primate populations and is easily accessible for research; conservation education, especially regarding hunting; and the development of alternative economic models for the local people to reduce the likelihood of them selling off their lands to logging companies.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9FFF9AFFF666F0F648FA46.taxon	materials_examined	Indonesia, Mount Sekumbang, south-east of Banjermasin, South Kalimantan.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9FFF9AFFF666F0F648FA46.taxon	discussion	P. rubicunda is closely related to P. melalophos, P. sumatrana, P. bicolor, and P. mitrata from Sumatra and P. comata from Java. There is much intergradation among subspecies, as well as variation within them in terms of overall body color. Moreover, there is an east-west division on the basis of the color of the hands and feet, and a north-south division between those with narrow and broad crests; the former have a U-shaped parting on the nape, while in the latter there is only an indistinct whorl on the nape. There is also an undescribed form from north-eastern Borneo (Sarawak, Sabah, and possibly Brunei). It is similar to the subspecies rubicunda, but with a broader crest. Five subspecies recognized.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9FFF9AFFF666F0F648FA46.taxon	distribution	Subspecies and Distribution. P. r. rubicundaS. Miller, 1838 — Borneo (SEKalimantan, roughlySoftheMahakamRiverandEoftheBaritoRiver). P. r. carimataeG. S. Miller, 1906 — KarimataI, offWBorneo. P. r. chryseaDavis, 1962 — NEBorneo, confinedtoaverysmallareainESabahStatenearKinabatangan. P. r. ignitaDollman, 1909 — Borneo, inSarawakState, fromtheBaramRiverattheborderswithBrunei, andinKalimantanStotheKapuasRiver; possiblyinBrunei. P. r. rubida Lyon, 1911 — Borneo (SW Kalimantan, roughly S of the Kapuas River and W of the Barito River).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9FFF9AFFF666F0F648FA46.taxon	description	Descriptive notes. Head — body 44 - 58 cm (males) and 48 - 52 cm (females), tail 63 — 80 cm (males) and 68 - 73 cm (females); weight 6 - 7 kg (males) and 5.5 - 6 kg (females). The Maroon Langur is generally wine-red to orange above and lighter yellowish or creamy on its belly, with black or brown hands and feet. There is a prominent, forwardsweeping black brow fringe and a frontal whorl behind, from which a large crown crest sweeps to the back of head that may be either narrow or broad. Facial skin is bluish-gray, exceptfor pinkish upperlip and chin. The “ Maroon Red Langur ” (Pr. rubicunda) is maroon to mahogany-red, with underparts and inner surfaces of limbs slightly paler. Crown crest is narrow. Hands and feet are washed with black, and tail has scattered black hairs, especially toward tip. P. r. rubida is like the Maroon Red Langur, but more yellowed; its hands and feet are only slightly blackened (or not at all), and its crown crest is narrow. The “ Orange-naped Red Langur ” (P. rn ignita) is foxy-red, with light colored hands and feet. Its crown crest is broad. The “ Orange-backed Red Langur ” (P. r. chrysea) is a rich reddish-gold above, and paler on abdomen and sides of thighs. Its digits are buff washed with brown, and its tail is washed with brown at tip. The “ Red-naped Red Langur ” (P. r. carimatae) is dark brick-red above, with somewhat darkened hands and feet. Its crown is tinged with gold and has a narrow crest.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9FFF9AFFF666F0F648FA46.taxon	biology_ecology	Habitat. Primary and secondary lowland rainforest and some dipterocarp, swamp, and riparian forest. The Maroon Langur occasionally enters plantations and gardens and is able to tolerate forests that have regenerated after they have been logged. Densities decline above elevations of 1200 m, and it is generally not found above 2000 m.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9FFF9AFFF666F0F648FA46.taxon	food_feeding	Food and Feeding. The Maroon Langur eats young leaves, seeds, fruits, mature leaves, and flowers. Termites and soil are also eaten on occasion. The Maroon Langur’s annual diet contains mostly young leaves (36 - 5 %), seeds (30 %), fruits (19 %), flowers (11 %), mature leaves (1 %), and insects and unidentified items (2: 5 %). Diets vary throughout the year as availabilities of food sources shift. In Sabah, a group relied on leaves and fruits of lianas for ¢. 32 % of their total diet; the rest came from various tree species. A single group exploited more than 102 plant species in a single year.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9FFF9AFFF666F0F648FA46.taxon	breeding	Breeding. There is no evident seasonality in births of Maroon Langurs. Neonates are entirely white. They are weaned at ¢. 15 months old. Females reach sexual maturity at c. 2 years old.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9FFF9AFFF666F0F648FA46.taxon	activity	Activity patterns. The Maroon Languris diurnal and almost entirely arboreal.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9FFF9AFFF666F0F648FA46.taxon	biology_ecology	Movements, Home range and Social organization. Social groups of Maroon Langurs contain 2 - 12 individuals, with one male, 2 — 4 females and their offspring. They also form all-male groups. Home ranges are 30 — c. 90 ha, and daily travel distances are 200 - 900 m. Male Maroon Langurs give loud calls that are short series of repeated “ ka-ka-kaka-ka ” notes that can be heard over several hundred meters. They are uttered regularly before dawn when the group is still at its sleeping site, sometimes during the day, and also in the evening when the group enters sleeping trees. They also give these calls when confronting neighboring groups.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9FFF9AFFF666F0F648FA46.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Least Concern on The [UCN Red List, including the subspecies rubicunda, rubida, and ignita, but the subspecies chrysea and carimatae are classified as Data Deficient. The Maroon Langur is protected in Sarawak and Sabah (Malaysia). It is still common and has a wide distribution. Habitat loss due to expanding oil palm plantations is a major threat, but the Maroon Langur seems to be adaptable and can occur in some secondary habitats. It is hunted forits meat and bezoars (a visceral mass believed to be an antidote to poison) in some areas. The subspecies rubida occurs in Betung Kerihun National Park, Bukit Baka-Bukit Raya National Park, Gunung Palung National Park, and Tanjung Puting National Park; the Maroon Red Langur occurs in Kutai National Park, Pleihari Martapura Nature Reserve, and Sungai Wain Protected Forest; and the Orange-naped Red Langur, or a related (undescribed) subspecies, occurs in Sungai Kayan Sungai Mentarang National Park — all in Kalimantan. The Orange-naped Red Langur also occurs in Sapagaya Forest Reserve and Danum Valley Conservation Area in Sabah, Malaysia. The Red-naped Red Langur is restricted to the 176 km? Karimata Island. About 50 % of the forest on the island has been lost, and the Red-naped Red Langur does not occur in any protected area, suggesting that it might be critically endangered. The Orange-backed Red Langur is only known from the type series. There are no protected areas in its putative distribution.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9FFF9BFAF56313FBCFFA6A.taxon	materials_examined	Malaya, Malacca.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9FFF9BFAF56313FBCFFA6A.taxon	discussion	In addition to the subspecies below, an extremely pale form, described as white with a little gray on its back and a blackish crest, has been reported from near Jambi in eastern Sumatra. The subspecies are rather poorly defined. Four subspecies recognized.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9FFF9BFAF56313FBCFFA6A.taxon	distribution	Subspecies and Distribution. P. s. siamensisS. Muller & Schlegel, 1841 — MalayPeninsula, inSThailand (populationisolatereportedat 6 ° 22 " N, 101 ° 08 E, wellwithinthedistributionofP. femoralisrobinsoni) andmostofPeninsularMalaysia (itisabsentintheNWtotheNtothePiahValleyinPerakState, andintheS). P. s. canaG. S. Miller, 1906 — CESumatra (betweentheSiakandIndragiririvers) andtheRiauArchipelago (PulauKundurI). P. s. paenulataChasen, 1940 — CESumatra (confinedtoasmallwedgeofcoastalforest, withapopulationisolatereportedfromnearLakeToba); itisseparatedfromP. femoralispercurabytheRokanRiver. P. s. rhionis G. S. Miller, 1903 — Riau Archipelago (for certain only from Bintan but may occur also on Batam and Galang Is).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9FFF9BFAF56313FBCFFA6A.taxon	description	Descriptive notes. Head-body 34 - 39 cm, tail 67 - 89 cm; weight 4.3 - 8.2 kg. The Palethighed Langur is similar to the Banded Langur (P. femoralis), but much paler grayishbrown in color and with a blackish brow, hands, and feet. Underside is whitish, continuous with a grayish-whitish zone on outsides of thighs. In general, it is pale brown-gray to buff above with a whitish underside (extending to inner surfaces of limbs and reaching down to wrists and ankles, and forward to throat and cheeks). Hands and feet are raven-black, asis tail, except for some white underits base. There is some slight clinal variation running north to south. The “ Bintan Pale-thighed Langur ” (FP. s. rhionis) 1 s yellow-brown, with a gray fringe above and a white underside, the latter color reaching ankles but not under tail. Its head and lower limbs are blacker. There is usually a single whorl on forehead. Its buccal depigmentation extends up to the nose. The “ Riau Pale-thighed Langur ” (P. s. cana) is similar to the Bintan Pale-thighed Langur, but it has a rather darker tail and limbs. Its fur is yellowish-gray above, with a darker head. Its white underside extends to chin, cheeks, wrists, and ankles, and backs of thighs have a wide gray area. Its tail is lighter below but not white. Its face is gray, with bluish eye-rings and a pinkish muzzle, and number of forehead whorls varies from two, one, or none. In the “ Mantled Pale-thighed Langur ” (P. s. paenulata), dorsal side is dark reddish-brown, while underside and inner surfaces of limbs are whitish to pale grayish-brown. Its outer limbs are brown, except for gray-white on backs of thighs and arms. Its hands and feet are black. Its tail is like its back above but paler below. Its face is black, with complete white eye-rings and pinkish lips, and its crest is small and black with a pale patch in front.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9FFF9BFAF56313FBCFFA6A.taxon	biology_ecology	Habitat. Primary and secondary rainforest, swamp, mangrove forest, and rubber tree plantations. Only a little information on habitat and ecology is available for the Palethighed Langur, but it is most likely similar to closely related species. The Bintan Palethighed Langur occurs in lowland wet forest, swamp forests, secondary regrowth, and “ rubber gardens ” (fallow secondary forest in slash-and-burn agriculture that is planted with rubber trees).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9FFF9BFAF56313FBCFFA6A.taxon	food_feeding	Food and Feeding. The Pale-thighed Langur eats mainly young leaves and fruits, along with seeds and flowers.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9FFF9BFAF56313FBCFFA6A.taxon	breeding	Breeding. There is no set breeding season for the Pale-thighed Langur. The female usually gives birth to a single young, although twins have been reported. The gestation period is c. 168 days. Infants are whitish-gray, with a distinctive dark cross pattern along their backs and shoulders.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9FFF9BFAF56313FBCFFA6A.taxon	activity	Activity patterns. The Pale-thighed Langur is diurnal and arboreal.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9FFF9BFAF56313FBCFFA6A.taxon	biology_ecology	Movements, Home range and Social organization. The Pale-thighed Langur has not been studied in the wild. Social groups have up to 18 individuals, with a single adult male, up to eight females, and their offspring.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9FFF9BFAF56313FBCFFA6A.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Near Threatened on The IUCN Red Lust, including the subspecies siamensis, cana and paenulata, but the subspecies rhionis is classified as Data Deficient. Deforestation and conversion of habitat most likely are major threats to the Pale-thighed Langur, and hunting to a lesser extent. Oil palm plantations are fast replacing lowland forests whereit lives. The “ Malayan Palethighed Langur ” (Ps. siamensis) occurs in Taman Negara National Park in the states of Pahang, Kelantan, and Terengganu, and Krau Wildlife Reserve in Pahang, Malaysia. It is not clear if the other subspecies occur in any protected area. Their taxonomy is disputed and distributional boundaries are unknown, so that an assessment ofits protected status is difficult. Further research on the Pale-thighed Langur is urgently needed to clarify conservation needs of the subspecies.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9EFF9BFF2B60CFFEF7F46A.taxon	materials_examined	Indonesia, Bunguran Island, North Natuna Islands.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9EFF9BFF2B60CFFEF7F46A.taxon	discussion	P. natunae has been variously considered a subspecies of P. femoralis, P. siamensis, and P. melalophos, but it is now regarded as a valid species. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9EFF9BFF2B60CFFEF7F46A.taxon	distribution	Distribution. Natuna Is (Buguran I), off the NW coast of Borneo.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9EFF9BFF2B60CFFEF7F46A.taxon	description	Descriptive notes. Head-body 43 - 61 cm, tail 61 - 84 cm; weight 5.9 - 8.2 kg. The Natuna Islands Langur is dark grayish-brown above, with head and lower parts of limbs blacker. White of underside extends to backs of thighs, chest, chin, and sometimes wrists and ankles, although not under tail. Cheek whiskers are exceptionally bushy and prominent, being mainly white except for their upper margins. Forehead whorl or whorls are absent or very indistinct. White eye-rings are very large, especially below eyes, but their upper and lower halves do not connect. Buccal depigmentation extends to the nose, with an additional, slightly depigmented area bordering frontal hair on, and lateral to, the glabella. Chest hairs are directed outward, except at the midline.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9EFF9BFF2B60CFFEF7F46A.taxon	biology_ecology	Habitat. Primary and logged forest and rubber tree plantations.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9EFF9BFF2B60CFFEF7F46A.taxon	food_feeding	Food and Feeding. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9EFF9BFF2B60CFFEF7F46A.taxon	breeding	Breeding. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9EFF9BFF2B60CFFEF7F46A.taxon	activity	Activity patterns. The Natuna Islands Langur is diurnal and arboreal.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9EFF9BFF2B60CFFEF7F46A.taxon	biology_ecology	Movements, Home range and Social organization. Social groups of the Natuna Islands Langur are small, averaging 3 - 5 individuals.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9EFF9BFF2B60CFFEF7F46A.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Vulnerable on The IUCN Red List. The Natuna Islands Langur is not protected under Indonesian law. It is threatened by forest loss and degradation (logging) and possibly offshore petroleum production and defense facilities. Natuna Islands Langurs are often kept as pets. There are no protected areas in its distribution. In 2002, the total population was estimated at less than 10,000 individuals in two subpopulations.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9EFF9BFF296ECFF88DF265.taxon	vernacular_names	Thomas's Langur	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9EFF9BFF296ECFF88DF265.taxon	materials_examined	Indonesia, Sumatra, Aceh, Langkat.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9EFF9BFF296ECFF88DF265.taxon	discussion	This species is monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9EFF9BFF296ECFF88DF265.taxon	distribution	Distribution. N Sumatra (Aceh Province), mostly N of the Wampu and Alas (= Simpang Kiri) rivers; a population has also been discovered S of the Alas at ¢. 3 ° 50 ” N.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9EFF9BFF296ECFF88DF265.taxon	description	Descriptive notes. Head-body c. 55 cm, tail 69 - 76 cm; weight 7 - 8 kg. There is no sexual dimorphism in size. Thomas’s Languris similar to the Javan Langur (P. comata), but differs slightly in having white patches above each eye and a vertical black stripe down the middle of its crest, with an additional black stripe on each side. Upperparts (including hands, feet, and face) are grayish-black, while underside is creamywhite with traces of yellow, the latter tone extending to inner surfaces of limbs, ventral surface of tail, and ears. Area surrounding muzzle is often pinkish.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9EFF9BFF296ECFF88DF265.taxon	biology_ecology	Habitat. Primary and secondary lowland rainforest, swamp forest, lowland alluvial (riparian) forest, and fruit and rubber tree plantations. The highest densities of Thomas’s Langurs have been recorded in swamp forest, with densities somewhat lower in lowland alluvial and lowland forests. They are also found in mosaics of rubber plantations and primary hill dipterocarp forest, and they tolerate selectively logged forest. Thomas’s Langur prefers the understory. They occur at elevations of 1500 - 2400 m and in lower parts of the subalpine zones up to 3400 m in the Leuser Ecosystem. Rainfall is high where they occur (3288 - 4575 mm / year), and forests are humid throughout the year. They were studied in a forest-plantation mosaic at Bohorok (1981 - 1984) and alluvial lowland rainforest at Ketambe Research Station, Aceh (1987 - 2001).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9EFF9BFF296ECFF88DF265.taxon	food_feeding	Food and Feeding. Thomas’s Langurs eat mainly young leaves, fruits and seeds, along with flowers, stems, toadstools, stalks of coconuts, soil, and animal prey (including insects and ground snails). Groups often raid crops. At Ketambe, Thomas’s Langur fed from 218 species of trees and lianas: 191 for fruits, 69 for leaves, and 28 for flowers. In forest-plantation mosaic at Bohorok, the number of species exploited was considerably reduced: 26 including cultivated banana (Musaceae) and seeds of the rubber tree (Hevea brasiliensis, Euphorbiaceae). Diets of groups of Thomas’s Langur at Ketambe comprised 36 - 1 % fruits and seeds, 30 - 8 %), young leaves, 13 - 4 % mature leaves, 3 - 6 % flowers, and 6 % animal prey (including ants and snails), with the remainder, soil from termite mounds, algae, and other food items. At Bohorok, they ate more fruits (more than 50 % of the diet) and leaves (¢. 36 %).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9EFF9BFF296ECFF88DF265.taxon	breeding	Breeding. Births of Thomas’s Langurs occur throughout the year. The natal coatis white, with a thick dark dorsal stripe. Females are considered adult at 60 months, about one year before the average age at which infants are born (74 months) and the youngest age at which a female in a study population gave birth. Interbirth interval averages 22 months (27 months counting only instances in which the previous infant survived). Females have a birth rate of c. 0 - 44 infants / year. Mortality is high in the first year oflife (48 % for male infants, 43 % for females). Instances of infanticide have been observed. In the wild, one female lived for more than 20 years.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9EFF9BFF296ECFF88DF265.taxon	activity	Activity patterns. Thomas's Langur is diurnal and largely arboreal. An activity budget recorded in one study was: 60 - 4 % of their day resting, 31 - 7 % feeding, 4 - 9 % traveling, and 3 % engaging in social activities. They sleep high up in tall trees that are clear of lianas.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9EFF9BFF296ECFF88DF265.taxon	biology_ecology	Movements, Home range and Social organization. Thomas’s Langurslive in unimale — multifemale groups of 2 - 16 individuals, and they also form all-male groups. Two-male groups were recorded at Bohorok, and they were always larger (9 - 21 individuals). Both males and females disperse from their natal groups, and females also transfer among groups. Sometimes all females leave a group and transfer to another, resulting in an all-male group being left behind. Sometimes females leave a group to join a solitary male. Home ranges averaged 27 ha at Ketambe, but they were smaller at Borohok at 14 ha. The largest home range was 53 ha (at Ketambe). Home ranges tend to be larger in groups that have been stable for some years. Newly established groups have small home ranges, independent of the group’s size. At Ketambe, male group tenure was five days to 72 months, with a median of 60 months. Males are typically resident in only one mixed-sex group; establishing their residency at c ¢. 7 years old and maintaining their status until they are c. 13 years old. Daily distance traveled was smaller at Bohorok (average 683 m / day) than at Ketambe (average 1068 m / day). Thomas’s Langurs have very distinct long calls, which distinguish them from all other langurs on Sumatra, especially neighboring Black-crested Sumatran Langur (P. melalophos). These calls show age, individualism, and contextual variation. Thomas’s Langurs utter different calls depending on whether or not other individuals are present and whether or not they give alarm calls. Loud calls are uttered in the morning before leaving their sleeping trees, during encounters with other groups, and as an alarm call after spotting a predator. Males give loud calls and chase each other when groups meet; females of each group generally ignore each other. The aggressive behavior is believed to result from the male’s defense of his females rather than defense of the territory itself.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FF9EFF9BFF296ECFF88DF265.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Vulnerable on The IUCN Red List. Thomas’s Langur is protected by Indonesian law. It is threatened mainly by habitat loss. Hunting does not seem to be a significant threat, but loss of primary habitat due to logging and conversion to oil palm plantations is probably displacing some populations. Thomas’s Langurs are sacred to the Batak people and consequently not hunted by them, although some marginal hunting occurs elsewhere in its distribution. Thomas’s Langurs occur in Gunung Leuser National Park, Sumatra.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA0FFA5FF2164BEF596F9CD.taxon	materials_examined	Vietnam, “ Cochin-China. ”	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA0FFA5FF2164BEF596F9CD.taxon	discussion	There is a small area of sympatry between P. nemaeus and P. cinerea in northern Quang Nam Province, Vietnam, and in north-eastern Cambodia (Virachey National Park), with some hybridization. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA0FFA5FF2164BEF596F9CD.taxon	distribution	Distribution. EC & SE Laos, N & C Vietnam (but very fragmented), and a small area in NE Cambodia (Voensei, Ratanakkiri Province). Old records from Hainan I appear to be erroneous.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA0FFA5FF2164BEF596F9CD.taxon	description	Descriptive notes. Head-body 55 - 63 cm (males) and 49 - 57 cm (females), tail 52 — 66 cm (males) and 42 - 59 cm (females); weight 811 - 6 kg (males) and 6 - 8 kg (females). Coloration of the Red-shanked Douc is variable. Males and females are the same and have the typical features found in the species of Pygathrix. Back, crown, and upper arms are dark gray-agouti. Chest, belly, and throat are very pale whitish-gray. Below the white throat, a broad orange collar is bordered by a black line underneath that joins black parts on shoulders. Back of hands and forearms, nearly to elbows, are white; hands and feet are black. Upper legs are black, lower legs are chestnut from knees to ankles, and maroon hairs cover feet. Face is surrounded by white whiskers as long as 12 cm in adult males. A wide forehead band is black. Above base of tail is a triangular patch of white fur, which in males is further adorned by a pair of tufts at each anterior angle. Facial skin is orange to yellow-brown, except around mouth and chin, where it is white. Tail is white and has a thin tassel. Penis is pink, and scrotum is white. In a small area in north-eastern Cambodia and in the southernmost part of Laos, coloration of the Red-shanked Douc resembles that of the Gray-shanked Doucs (P. cinerea). White color on forearms is highly variable and in some individuals is reduced to small patches on wrists or completely missing. Forearms are gray-agouti. Maroon color on lower legs is also reduced to a reddish shine on gray-agouti legs. Natal coloration of the Red-shanked Douc is characterized by a light gray-agouti stripe on the back along the spine and white on the belly that extends over sides of the body to the back and to arms. Legs are light chestnut. From the forehead up to the crown, hairs are light chestnut; back of head is gray-agouti. Tail is whitish to grayish-white and has a thin tassel. There is a considerable variation in facial coloration, from a complete dark slate-gray face to more or less extended yellow patches under and over eyes, and to wider yellow eye-rings with only restricted gray parts around mouth, nose, and forehead. During the first five months oflife, forearms change to white, upper legs change to dark gray, and lower legs change to darker maroon. Chestnut color on head changes to gray, with a darker gray or blackish band over the forehead. Tail color changes to white. Slategray parts on face become paler over a long period of time, c. 3 years, changing to the yellow-orange facial skin of adults.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA0FFA5FF2164BEF596F9CD.taxon	biology_ecology	Habitat. Subtropical and tropical evergreen or semi-deciduous broadleaf forest, (classified as lowland and montane forest), karst formations, and occasionally mixed bamboo forest. Red-shanked Doucs are observed from sea level up to an elevation of 1600 m. They prefer high forest canopies but occasionally go to the ground for water or to eat minerals.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA0FFA5FF2164BEF596F9CD.taxon	food_feeding	Food and Feeding. The Red-shanked Douc is largely folivorous but also eats fruits and seeds. Its diet consists of ¢. 80 % leaves. Stomachs of 13 individuals contained 63 % leaves and buds, along with flowers, fruits, and seeds. In Vietnam, 125 species of plants have been identified in the diet. Leaves are eaten from 83 species, fruits from 79, bark from five, and roots from one species. Young leaves that are high in crude protein and low in fiber are preferred. On average, leaf intake per individual is 1700 g / day.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA0FFA5FF2164BEF596F9CD.taxon	breeding	Breeding. Male Red-shanked Doucs reach sexual maturity at 5 - 6 years old and females at 3 - 5 - 5 years. Females have 28 - 30 day reproductive cycles, and their normally white genital area becomes suffused with red when sexually receptive and during pregnancy. A single young is born after a gestation of 180 - 190 days. Young are weaned at 9 - 16 months old, varying depending on whether or not the female has a new pregnancy. Interbirth intervals are 9 - 26 months. Infants are gray and ginger, with a mostly black face (mottled with pale pink). Allomothering appears to be common. Individuals have lived up to 25 years in captivity.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA0FFA5FF2164BEF596F9CD.taxon	activity	Activity patterns. Red-shanked Doucs are diurnal and arboreal. There are feeding peaks in the morning and afternoon, with a long rest at midday. They spend ¢. 44 % of their day feeding (30 % on leaves and 14 % on fruits), 15 % engaging in social activities including grooming and playing, 14 % inactive, 12 % resting, 10 % traveling, and 5 % engaging in other activities. The time budget of daily activities varies among groups with and without human disturbance. A disturbed group rests less (c. 5 % less) and travels more. Leaping and jumping are the most common form of locomotion; brachiating may also be used when traveling in the forest canopy, particularly if individuals are fleeing or immediately before they begin leaping. Arboreal quadrupedalism is another mode of travel, which is often used where trees have open branches.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA0FFA5FF2164BEF596F9CD.taxon	biology_ecology	Movements, Home range and Social organization. Group structure of the Red-shanked Douc is either unimale — multifemale or multimale — multifemale, with 3 - 20 individuals / group (5 - 10 most common). The male-female ratio averages 1: 2 - 4. Group composition estimated from several populations is 21 % adult males, 51 % adult females, 18 % subadults, and 10 % infants. Several groups sometimes travel and forage together for a while, forming bands of 50 or more individuals. A low-pitched growl is given as a threat, and there is also a short, harsh distress squeal. There is a specific “ play face, ” in which eyes are closed (exposing very blue eyelids). The characteristic fixed stare is actually a threat display, with an open-mouthed grimace being the corresponding gesture of submission, which is also used to initiate grooming. There is a pre-mating sexual signal given by both sexes, which consists of placing the jaw forward, raising eyebrows and then lowering them, and then shaking the head.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA0FFA5FF2164BEF596F9CD.taxon	conservation	Status and Conservation. CITES Appendix I. Classified as Endangered on The IUCN Red List. The Red-shanked Douc is protected in all the three countries where it occurs. The population in Laos is probably quite large and is, as such, the most important for Red-shanked Doucs. The largest populations occur in Nakai-Nam Theun and Hin Namno national biodiversity conservation areas. In Vietnam, it has declined significantly in recent decades because of hunting, logging, and the conversion ofits forests for coffee, rubber, and cashew plantations. Hunting is the major threat to Redshanked Douc, most often for subsistence and traditional Asian medicine. The population is highly fragmented. The largest population in Vietnam exists most probably in Phong Nha-Ke Bang National Park, with an estimated 1300 individuals. Smaller populations in Vietnam exist in Pu Mat Nature Reserve and Bach Ma National Park (Nghe An and Thua Thien Hue provinces) and Son Tra Nature Reserve (Danang Province).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA0FFA6FA2C6365FB03F46C.taxon	materials_examined	Vietnam, Gia Lai Province, Play Ku.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA0FFA6FA2C6365FB03F46C.taxon	discussion	Play Ku is a wildlife trade center, and P. cinerea does not occur there naturally. The holotype’s origin is most probably Kon Ka Kinh National Park. There is a small area of sympatry with P. nemaeus in northern Quang Nam Province, Vietnam, and in north-eastern Cambodia (Virachey National Park), with some hybridization. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA0FFA6FA2C6365FB03F46C.taxon	distribution	Distribution. C Vietnam between 16 ° N and 13 ° 38 " N (Quang Nam, Kon Tum, Quang Ngai, Gia Lai, and Binh Dinh provinces), and a small part in NE Cambodia.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA0FFA6FA2C6365FB03F46C.taxon	description	Descriptive notes. Head-body 55 - 66 cm (males) and 51 - 568 cm (females), tail 56 — 65 cm (males) and 54 - 62 cm (females); weight 8 - 12.5 kg (males) and 4.7 - 11.7 kg (females). Males and females have the same pelage color. Back, nape, and crown of the Gray-shanked Douc have a gray-agouti color that extends to arms, except for insides of forearms. Chest, belly, and throat are very pale whitish-gray. Below its white throat, a broad orange collar is bordered underneath by a black line that joins black parts on shoulders and down to inside upper arms and forearms. Color of legs varies from light gray-agouti to very dark gray-agouti, nearly black, and becoming black on inner side of thighs. Hands and feet are black. Tail is white and has a thin tassel. Above its tail root is a triangular patch of white fur, which in males is further adorned by a pair of tufts at each anterior angle. Facial skin is orange to yellow-brown, except around mouth and chin, where it is white. Eyes are slanted at c. 14 - 5 ° from horizontal. Long white cheek whiskers frame face to the sides and curve downward, much longer in males than females. Gray-agouti hairs on head have a cape-like form on nape, and a small black frontal band occurs over forehead. Newborn Gray-shanked Doucs have a pale gray-agouti back, a white belly, and very light gray arms. Inner sides of legs are white, and outer sides are light chestnut. From forehead up to the crown, hairs are light chestnut; back of head is gray-agouti. Tail is whitish to grayish-white and has a thin tassel. There is wide variation in facial coloration, from a dark slaty-gray face with more or less extended yellow patches under and over eyes to wider yellow eye-rings with only restricted gray parts around mouth, nose, and forehead. During the first three months of life, young lose the reddish tinge on their heads and legs and change to gray. Tail color changes to white. Slaty-gray parts of face become pale over about the next three years, changing to the yellow-orange facial skin of adults.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA0FFA6FA2C6365FB03F46C.taxon	biology_ecology	Habitat. Evergreen lower and montane moist subtropical forest and mixed broadleaf and needle-leaf moist subtropical forest, even in degraded parts, from elevations of c. 200 m up to c. 1500 m. Winter and spring are dry and cool, and summer and autumn are humid and warm. Mean annual temperature is ¢. 22 ° C, mean daily maximum temperature in the warmest month is ¢. 32 ° C, mean daily minimum temperature in the coldest month is ¢. 11 ° C, and absolute temperature minimum is ¢. 8 ° C.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA0FFA6FA2C6365FB03F46C.taxon	food_feeding	Food and Feeding. Gray-shanked Doucs are folivorous but also include fruits and seeds in their diets. Over the year, young leaves are preferred (c. 50 % of the diet), with mature leaves contributing ¢. 10 %, ripe fruits 22 %, and unripe fruits 10 %. There is high seasonal turnover of food items selected. In the dry season, young leaves comprise ¢. 80 % of the diet, but in the wet season, ¢. 18 %. The proportion of fruits in the diet changes from less than 10 % in the dry season to ¢. 35 % in the wet season. In all, 166 tree and plant species from 40 families have been identified as providing food for Gray-shanked Doucs. They have a rather high demand for water during digestion, and they use water sources on the ground. About 60 % of the water demand is satisfied from their food, and 40 % from drinking. Water consumption is correlated with air temperature and increases from 0 - 10 ml / g body weight at 24 ° C to 0 - 16 ml / g body weight at 32 ° C.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA0FFA6FA2C6365FB03F46C.taxon	breeding	Breeding. Births of Gray-shanked Doucs occur throughout the year, with a peak in January-March. They are sexually receptive every 28 - 30 days. The gestation period is 165 - 190 days. A single young is born.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA0FFA6FA2C6365FB03F46C.taxon	activity	Activity patterns. Gray-shanked Doucs are diurnal and largely arboreal, but also terrestrial, going to the ground to drink water and lick minerals. Their daily activities begin at 05: 00 - 06: 00 h and cease at 17: 00 - 18: 45 h. Activity budgets change seasonally. In the wet season, they spend, on average, 41 % oftheir time resting, 11 % feeding, 22 % moving, and 26 % socializing. In the dry season, c. 32 % of their time is spent resting, 13 % feeding, 31 % traveling, and 24 % socializing, which also includes interactions among adult males. Activity budgets vary significantly between males and females and age classes. Females spend more time feeding (14 %), resting (42 %), and moving (20 %) than males (11 %, 37 %, and 17 %). Males, on the other hand, spend more time engaged in social activities (35 %) than females (24 %). Subadults andjuveniles spend less time feeding (c. 7 %) and more time engaging in social activities (¢. 28 %) than either adult males or females. Gray-shanked Doucs are largely quadrupedal but frequently brachiate when traveling in the forest canopy.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA0FFA6FA2C6365FB03F46C.taxon	biology_ecology	Movements, Home range and Social organization. Daily travel distance of Grayshanked Doucs is correlated with group size; smaller groups travel less than larger groups. Groups with less than ten individuals travel, on average, less than 500 m / day, with a minimum of 50 m / day for a group of six. Groups with more than 20 individuals travel, on average, more than 2000 m / day, with a maximum of 4000 m / day for a group of 29. Thereis little information on home range size. A large band, consisting of six one-male groups that sometimes traveled together, used a home range of 984 ha. Unimale — multifemale groups in Kon Ka Kinh National Park, Vietnam, averaged 14 - 8 individuals. Large bands of up to 88 individuals result from up to six groups merging in a fission-fusion system. The most common one-male group had 6 - 9 individuals (30 %), followed by 2 - 5 individuals (23 %) and 10 - 15 individuals (21 %, n = 179). Grayshanked Doucs have four different social units: unimale-multifemale group, all-male groups, multimale-multifemale groups, and single males. The unimale-multifemale group is the most common. A male-to-female sex ratio of c. 1: 2 (216 individuals in 24 groups) and age class composition of c. 65 % adults, 18 % subadults, 11 % juveniles, and 4 % infants have been recorded. Both sexes disperse. The population density in Kon Ka Kinh National Park is estimated at 0 - 6 ind / km?.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA0FFA6FA2C6365FB03F46C.taxon	conservation	Status and Conservation. CITES Appendix I. Classified as Critically Endangered on The IUCN Red List. Since 2000, the Gray-shanked Douc has been listed as one of the “ World’s 25 Most Endangered Primates. ” It is protected by law in Vietnam and Cambodia. It was first discovered in 1995 when an individual was confiscated from a restaurant in northern Vietnam. The total population is estimated to be less than 1000 individuals, occurring in a number of isolated populations. The largest population, numbering ¢. 200 individuals, lives in Kon Ka Kinh National Park, in Song Thanh, Kon Plong and Kon Cha Rang nature reserves, and in unprotected areas in South Quang Nam Province. They also occur in Mom Ray National Park, Ngoc Linh and A Yun Pa nature reserves, and Ba To Cultural and Historical Site in Vietnam. Gray-shanked Doucs are threatened from habitat loss and poaching, both for food and traditional medicine. They are also susceptible to snaring when on the ground, especially in the more degraded habitats.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA3FFA7FFEB6EC8FB44FD39.taxon	materials_examined	Vietnam, Saigon.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA3FFA7FFEB6EC8FB44FD39.taxon	discussion	P. migripes is known to hybridize narrowly with P. nemaeus where their distributions overlap in northern Quang Nam Province, Vietnam, and in north-eastern Cambodia (Virachey National Park). Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA3FFA7FFEB6EC8FB44FD39.taxon	distribution	Distribution. E Cambodia (E of the Mekong River and S of the Srepok River) and SW Vietnam from c. 14 ° N to S Cat Tien National Park. The presence of this species in S Laos is suspected but has never been confirmed.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA3FFA7FFEB6EC8FB44FD39.taxon	description	Descriptive notes. Head-body 55 - 63 cm (males) and 54 - 60 cm (females), tail 69 - 84 cm (males) and 65 - 80 cm (females); weight 8 - 11 kg (males) and 6.8 - 2 kg (females). Males and females have the same fur coloration. Back, head, and arms of the Black-shanked Douc are dark-gray agouti, much darker than in the Redshanked Douc (P. nemaeus). Chest and belly are very light gray, nearly white. Upper and lowerlegs, feet, and hands are black. A white throat is surrounded by a chestnut collar that comes close to a black band running from shoulder to shoulder. Tail is white and has a thin tassel, and is longer than those of the other two doucs. Above tail root is a white triangular patch which in males is further adorned by a pair of tufts at each anterior angle. Face is bluish, with large yellow eye-rings. A black forehead band runs along sides of head down to shoulders. Ears are inside a black triangular area. Whiskers are short and thin, but longer hairs form a goatee. Penis is red, and scrotum is blue. A few individuals differ in coloration, e. g. white markings on wrists to completely white forearms, or more or less maroon lower legs, resembling the Red-shanked Douc, but with the typical and very different face coloration of the Black-shanked Douc. Newborn Black-shanked Doucs have a dark gray-agouti stripe on their back along the spine. White from the belly extends over side of the body to their back. Legs are chestnut, upper arms white, forearms light gray with a chestnut shine. Hands and feet are slate-gray. Crown is dark gray, with a chestnut band over the forehead. A very dark slate-gray marking on face runs from forehead over nose to mouth and chin, in contrast to wide yellow eye-rings that are not completely closed on nose. Sides of head, in front of ears, are white; ears are flesh-colored.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA3FFA7FFEB6EC8FB44FD39.taxon	biology_ecology	Habitat. Evergreen forest to other habitat types, extensively in semi-evergreen and mixed deciduous forest, and also dry deciduous dipterocarp forest, mixed bamboo — broadleaf forest, and coastal dry forest. Black-shanked Doucs are observed from sea level up to ¢. 800 m.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA3FFA7FFEB6EC8FB44FD39.taxon	food_feeding	Food and Feeding. The diet of Black-shanked Doucs depends on the forest type. It is very diverse, with no particularly favored species. Up to 152 plant species are exploited in southern Vietnam, including 117 for leaves, 42 for fruits, eleven for seeds, and one for its petioles. The 20 species eaten most often account for only ¢. 2 % of the diet. In dryer areas and at higher elevations in southern Vietnam, the diet comprises ¢. 55 % leaves, 30 % seeds and fruits, and 15 % flowers. In Seima Biodiversity Conservation Area, east Cambodia, percentages are 40 % leaves, 40 % seeds, ¢. 10 % fruits, and 10 % flowers. There is a significant association between time of day and food items selected for consumption. Seeds are exploited at the expense of all other food items in the early morning. Preferences change from leaves at midday to fruits in the afternoon. Males eat more seeds and fewer leaves than females. In southern Vietnam, relative amounts of leaves and fruits between morning and afternoon are nearly equal. There are differences in diets between dry and wet seasons, with 60 % leaves, 20 % flowers, and 17 % fruits (3 % other) in the dry season compared to 52 % leaves, 14 % flowers, and 34 % fruits in the wet season.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA3FFA7FFEB6EC8FB44FD39.taxon	breeding	Breeding. Offspring of Black-shanked Douc appear to be born throughout the year.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA3FFA7FFEB6EC8FB44FD39.taxon	activity	Activity patterns. The Black-shanked Douc is diurnal and arboreal. They become active shortly before sunrise, ¢. 05: 30 h, and begin the day with intense bouts of feeding that decrease later in the morning and are followed by resting, predominantly from 10: 00 h until 14: 00 h. There is a second period of activity in late afternoon. After 18: 00 h, activities cease and individuals prepare for sleep. The daily activity budget of a group, including both sexes and all ages except infants, averages 61 % resting, 27 % feeding, 7 % traveling, 3 % engaging in social behavior, and 2 % engaging in other activities. During the wet season, more time is spent resting (62 %) and less feeding (23 %) compared to the dry season with 55 % and 31 %. Black-shanked Doucs occasionally go to the ground to move or drink water. In one coastal population in southern Vietnam, groups spend up to 20 % of the day on large granite boulders. Quadrupedal locomotion dominates their movements (61 %), followed by leaping (17 %), brachiating (10 %), climbing (8 %), and dropping (free-falling) to lower levels of the canopy (4 %).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA3FFA7FFEB6EC8FB44FD39.taxon	biology_ecology	Movements, Home range and Social organization. There is little information about home ranges of Black-shanked Doucs. One study indicated they are c. 20 ha, but most probably larger, up to 30 - 50 ha. Daily travel distances are 700 - 1200 m. Groups in lowquality habitat travel farther. Large trees are used as sleeping sites, sometimes close to rivers. Groups are typically unimale-multifemale. Their size differs in different habitats. In two protected areas in South Vietnam, groups averaged 13 individuals (range 3 - 22) and ten individuals (5 - 15). In east Cambodia, average group size is 7 - 5 individuals. Males outside these groups are solitary or live in all-male groups. Groups come together in a fission-fusion behavior forming larger bands of up to 50 individuals. These bands might spread out over a diameter of ¢. 100 m and move as if they are one group. An all-male group has been seen with at least 17 individuals. In the Seima Biodiversity Conservation Area of Cambodia, the population density of Black-shanked Doucsis 1 - 2 groups / km?.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA3FFA7FFEB6EC8FB44FD39.taxon	conservation	Status and Conservation. CITES Appendix I. Classified as Endangered on The IUCN Red List. The Black-shanked Douc is protected by law in Cambodia and Vietnam. Although once thought to be rare with a limited distribution, or even extinct, recent fieldwork has demonstrated that populations are quite widespread and found in a variety of habitats. There are few data available from Cambodia, but one robust population estimate from Seima Biodiversity Conservation Area was ¢. 6000 groups, or a population of ¢. 42,000 individuals. Populations in Vietnam have suffered a massive decline, and are now very fragmented. Probably one of the largest populationsis that in Nui Chua National Park, Ninh Thuan Province, with 500 - 700 individuals; other populations occur in Cat Tien National Park in Dong Nai Province, Bi Dup Nui Ba Nature Reserve in Lam Dong Province, and in areas that are contiguous with protected areas in Cambodia. Hunting, mainly for traditional medicine, is the main threat. Destruction of natural habitatis also a threat; a large portion in central and southern Vietnam suffers from conversion of forest to farmland. Logging and clear-cutting for coffee, rubber, and cashew plantations have reduced natural habitats of the Black-shanked Douc. Most forests at low elevations have been cleared, and very little forest remains undisturbed.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA2FFA7FF2E66C3F73EFA4E.taxon	materials_examined	Yen Bay, Songkoi River, Vietnam.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA2FFA7FF2E66C3F73EFA4E.taxon	discussion	This species is monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA2FFA7FF2E66C3F73EFA4E.taxon	distribution	Distribution. NW Vietnam, known only from small forest patches in Ha Giang, Tuyen Quang, Bac Kan, and Thai Nguyen provinces.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA2FFA7FF2E66C3F73EFA4E.taxon	description	Descriptive notes. Head — body 55 - 65 cm (males) and 50 - 54 cm (females), tail 82 — 92 cm (males) and 65 - 73 cm (females); weight 14 kg (males) and 8 - 5 kg (females). The Tonkin Snub-nosed Monkey is smaller, darker, and more slender in build than the other four species of Rhinopithecus. It has more elongated digits. Males and females have the same pelage coloration. Back and outsides of legs and arms are dark brown. Head, belly, and inner sides of limbs are creamy-white, this latter tone extending high up flanks, around face, and down limbs (leaving only a thin black stripe down outer sides to hands and feet); on elbows, in particular, the creamy-white color from insides of arms nearly encloses outsides of arms. Hands and feet are black. There is a small orange collar on the throat. Facial skin is bluish-white around eyes and nose, and lips are pink and enlarged. Adults have a notably upturned nose. Ears with creamy fur stand out at right angles from head. Tail is very long (nearly 170 % of head — body length), with curly, intermixed black and strawcolored hairs along upper surface, a white underside, and a creamy-white tip with a thin tassel. Penis is black, and scrotum is white.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA2FFA7FF2E66C3F73EFA4E.taxon	biology_ecology	Habitat. Steep limestone karst hills with subtropical evergreen limestone forest and forest patches with bamboo at elevations of 200 - 1300 m.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA2FFA7FF2E66C3F73EFA4E.taxon	food_feeding	Food and Feeding. The Tonkin Snub-nosed Monkeyis a selective feeder. When feeding, it dedicates c. 40 % of its time to young leaves, 18 % to unripe fruits, 14 % to flowers, 10 % to young stems, and 6 % each to buds, seeds, and mature leaves. It feeds on young leaves, unripe fruits, and seeds of as many as 52 plant species. In the most recentfield study, proportions of young leaves in diets varied from 11 % in December — May up to 46 % in September, while proportions offruit fell from 47 - 63 % to 24 %. In December — May, 26 % of the diet in this study consisted of petioles, and the rest was flowers, mature leaves, seeds, and leaf buds.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA2FFA7FF2E66C3F73EFA4E.taxon	breeding	Breeding. Little is known about breeding of the Tonkin Snub-nosed Monkey. Births occur most during the first half of the year. A single young is born, with a creamyyellowish fur. After 1 - 2 months, the colorstarts to darken and changes slowly to brown.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA2FFA7FF2E66C3F73EFA4E.taxon	activity	Activity patterns. Tonkin Snub-nosed Monkeys are diurnal, arboreal, and occasionally terrestrial. They spend ¢. 33 % of their day resting, 20 % traveling, 30 % watching and feeding, and the rest engaging in different social activities. Groups do not have fixed sleeping sites. They generally sleep close to the ground, on steep mountainsides, away from the wind, at ¢. 5 - 10 m high in trees, sometimes in dense foliage. The Tonkin Snub-nosed Monkey travels on large tree branches by quadrupedal and occasionally bipedal walking and in the canopy, by leaping, jumping, and brachiating.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA2FFA7FF2E66C3F73EFA4E.taxon	biology_ecology	Movements, Home range and Social organization. Typical social unit of the Tonkin Snub-nosed Monkey is a unimale — multifemale group of 12 - 15 individuals, sometimes up to two dozen. The home range of one group was 380 - 560 ha. In the much depleted remaining population in Tat Ke and Khau Ca, the size of one-male groups was 7 - 13 individuals, with 2 - 6 adult females, 1 - 2 subadults, 1 - 3 juveniles, and 1 - 4 infants. Home ranges of the groups overlap considerably. Groups often travel, forage, and rest together in large semi-cohesive bands. Males disperse from their natal groups to form all-male groups; all-male groups in remnant populations consist of 1 - 2 adult males and 2 - 3 juvenile males. At times (it is not quite clear in what seasons or under what stimulus), these groups join to form large bands. In Khau Ca, a band of 78 - 81 individuals appeared to contain the entire population.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA2FFA7FF2E66C3F73EFA4E.taxon	conservation	Status and Conservation. CITES Appendix I. Classified as Critically Endangered on The IUCN Red List. The Tonkin Snub-nosed Monkey is protected by law in Vietnam. It has been on the list of the “ World’s 25 Most Endangered Primates ” since it wasfirst drawn up in 2000. Historically limited to areas east of the Red River, its distribution has been drastically reduced in recent decades because of massive deforestation and intensive hunting. It was thought to be extinct until its rediscovery near the town of Na Hang in 1989. Only a few degraded small populations exist. There are no exact numbers on the sizes of the subpopulations. An estimate is ¢. 30 individuals for Na Hang Nature Reserve. A population of 70 individuals was estimated for Cham Chu Nature Reserve but recent surveys failed to produce evidence that Tonkin Snub-nosed Monkey still exist in the area. About 100 individuals live in the Tonkin Snub-nosed Monkey Conservation Area in Khau Ca, Ha Giang Province, and probably 50 in Quan Ba District, close to the Chinese border in the same province. The total population is believed to be less than 200 individuals. Despite their legal protection and the fact that they are one of the world’s rarest primates, Tonkin Snub-nosed Monkeys continue to be hunted mercilessly for use in Chinese medicine; they are rarely eaten because the flesh is considered foul tasting. Illegal logging takes place in the habitat of the Tonkin Snub-nosed Monkey. Shifting and settled cultivation, as well as other land development activities, also pose threats. In the past, intense and unsustainable legal and illegal logging and gold mining were the biggest threats. Recently, development of a hydroelectric power project along the Gam River in Na Hang has led to an increase in the human population and the demand for meat (despite the poor taste), and thus increased hunting pressure. The total population was estimated at ¢. 250 in 2006, although this figure may now be higher due to possible occurrence in other areas. The Tonkin Snub-nosed Monkey occurs in four protected areas: Cham Chu Nature Reserve, Than Sa — Phuong Hong Nature Reserve, Tat Ke Sector of Na Hang Nature Reserve, and Khau Ca Habitat in Vietnam. It may also survive in Ba Be National Park, although the most recent search by Thanh Hai MaDong did not find any there; there may be a very small population in South Xuan Le Habitat.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA2FFA0FA2B60ECFB37F713.taxon	materials_examined	China, Yunnan, left bank of upper Mekong, Kiape, a day’s journey south of Atuntze (28 ° 25 ’ N, 98 ° 55 ’ E).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA2FFA0FA2B60ECFB37F713.taxon	discussion	This species is monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA2FFA0FA2B60ECFB37F713.taxon	distribution	Distribution. SW China, in SE Xizang Autonomous Region (= Tibet) and NW Yunnan Province (fragmented populations in the Yun Ling Mts), W of the Yangtze River and E of the Mekong River.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA2FFA0FA2B60ECFB37F713.taxon	description	Descriptive notes. Head-body ¢. 83 cm (males) and 74 - 83 cm (females), tail 52 - 75 cm; weight 15 - 17 kg (males) and 9 - 2 — 12 kg (females). Adult Yunnan Snub-nosed Monkeys are blackish-gray above with contrastingly white underparts, white extending high up on flanks, backs of thighs, and sides of neck to form a ring around face. Tail is black, with a cow-like tuft at the end, and hairs on backs of thighs are very long and wavy, especially in adult males. Brow is black, and there is a thin, high, forward-drooping crest on crown. Facial skin is bright pink or red on muzzle, with traces of pale yellowish or greenish around eyes; fur wedges separating these zones are not as striking as in the Golden Snub-nosed Monkey (R. roxellana). Female Yunnan Snub-nosed Monkeys have shorter fur than males, and pelage is much more contrastingly colored in adult males; white backs of thighs particularly stand out in the field.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA2FFA0FA2B60ECFB37F713.taxon	biology_ecology	Habitat. Temperate montane evergreen, broadleaf deciduous, and conifer forest, preferring fir-larch or cypress forests, also areas of bamboo. The Yunnan Snub-nosed Monkeylives at higher elevations (3000 - 4700 m) than any other non-human primate, where temperatures average below freezing for several months of the year and snow can accumulate to over a meter in depth. The region itself is snow-covered for at least half the year. Yunnan Snub-nosed Monkeys use lower elevations in winter than summer. It is sympatric with the Rhesus Macaque (Macaca mulatta).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA2FFA0FA2B60ECFB37F713.taxon	food_feeding	Food and Feeding. The Yunnan Snub-nosed Monkey is mainly folivorous and eats leaves and lichens, along with fruits, bark, berries, grasses, nuts, moss, seeds, shoots, and acorns. Lichens are important food items, making up ¢. 67 % of the diet. In Jinsichang, bamboo leaves form a large part of the diet. They will eat bird nestlings and also clay (geophagy). Diet varies largely by season.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA2FFA0FA2B60ECFB37F713.taxon	breeding	Breeding. Mating of the Yunnan Snub-nosed Monkey peaks in August-September, and most births occur in March — - May. A single young is born after a gestation of 189 - 198 days. In captivity, the birth interval is c. 2 years. Groups have a relatively high ratio of adult females to infants, indicating a low birth rate or low infant survival. Infant mortality in theirfirst harsh winter is 55 - 60 %. Newborns are completely white, except for a blue face and pink fingers and toes. Their fur turns to light yellow as juveniles and then to gray. In subadults (age 4 - 8 years), back, sides, and ends of limbs first turn to light brown and then eventually black. Sexual maturity occurs at 5 - 6 years for males and 4 - 5 years for females.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA2FFA0FA2B60ECFB37F713.taxon	activity	Activity patterns. The Yunnan Snub-nosed Monkey is diurnal, arboreal, and terrestrial. Most of the day is dedicated to feeding (39 %). It rests for c ¢. 35 % of the day, indulges in social activities for 16 %, and travels for c. 10 %. Yunnan Snub-nosed Monkeys sleep in dark conifer forest far from valley bottoms and often use the same sleeping site on consecutive nights.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA2FFA0FA2B60ECFB37F713.taxon	biology_ecology	Movements, Home range and Social organization. The Yunnan Snub-nosed Monkey lives in multimale-multifemale bands, comprised of unimale-multifemale and allmale groups. One band had 15 - 18 all-male groups. Large all-male groups may have as many as 14 - 16 individuals. Band sizes are ¢. 24 - 370 individuals, and they have home ranges of 1000 - 4000 ha, with overlap with neighboring groups of ¢. 30 %. Mean daily travel distance is ¢. 1300 m, and it is shorter in winter than summer. Indicating a matrilineal society, females tend to groom each other more than they groom males. It is probable that males rather than females disperse from their natal groups. Males often stand (or sit) guard over groups of up to eight playing infants and juveniles, while females forage and feed elsewhere. In captive groups, adult males can be protective of infants and young juveniles by intervening when older juveniles threaten or bully younger ones. Fission into smaller groupsis rare and usually only lasts for a day. Groups will split off from the band occasionally to drink in ponds, especially in the dry season. Males in all-male groups sometimes show aggression to other all-male groups at valued sleeping trees and waterholes and sometimes in feeding trees.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA2FFA0FA2B60ECFB37F713.taxon	conservation	Status and Conservation. CITES Appendix I. Classified as Endangered on The IUCN Red List. Illegal hunting and trapping of the Yunnan Snub-nosed Monkey is the major threat; they are often caught in snares set for other animals (e. g. chevrotains, Moschus). Habitat loss, mainly from logging, but also clearing for summer grazing and agricultural land, is also a concern. Pesticide spraying was the cause of extinction of one entire subpopulation of ¢. 50 individuals. In 2006, the total world population was estimated at less than 2000 individuals, with less than 1000 mature individuals in 15 subpopulations. Three subpopulations have been extirpated since 1994, but it is possible that still others await discovery. Eleven subpopulations occur in protected areas and four groups occur in unprotected forest fragments. The Yunnan Snub-nosed Monkey occurs in Hongla Snow Mountain National Nature Reserve, Baima Snow Mountain National Nature Reserve, and Laojun Mountain National Nature Reserve in China. Additional survey work is needed to find other subpopulations in non-surveyed regions. There is a major focus in China on captive breeding of the Yunnan Snub-nosed Monkey, with breeding pairs at the Kunming Zoo and Kunming Institute of Zoology. Most of these individuals were captured in the wild, and so far the program has achieved little success in being sustainable. The Yunnan Snub-nosed Monkey haslikely declined by well over 20 % in the last two generations (c. 25 years). A preliminary population viability analysis found that the five smallest subpopulations are at risk of declines and extinction in the next 100 years from effects of inbreeding and poaching, while the five largest subpopulations are apparently more secure. Since 1999, when a ban stopped most commercial logging in the region, habitat loss has slowed, butit could still be a major threat in the future. Clearing offorest land for summer grazing pasture reduced suitable habitat by 31 % between 1958 and 1997. Fires set for agriculture are a threat in some areas, particularly in the Tibetan Autonomous Region.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA5FFA0FFF36C27F5E2F89B.taxon	materials_examined	China, Van Gin Shan (= Fanjingshan), Guizhou.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA5FFA0FFF36C27F5E2F89B.taxon	discussion	This species is monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA5FFA0FFF36C27F5E2F89B.taxon	distribution	Distribution. SC China, restricted to Fanjingshan in the Wuling Mts, Guizhou Province.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA5FFA0FFF36C27F5E2F89B.taxon	description	Descriptive notes. Head — body 66 - 72 cm, tail 97 cm; weight 14 - 15 kg (males) and c. 8 kg (females). The Guizhou Snubnosed Monkey. is dark brown above, blending to a grayish color on its lower back, with underside and inner surfaces of limbs bright gingery-red in adult males (becoming overlain with black on chest and belly). Hands and feet are blackish. Face is bare and white, with black cheeks and dark bluish skin around eyes. It lacks cheek wedges seen in other species of Rhinopithecus. Brow is gingery-red, and ears are tipped with white. Tail is long (nearly 150 % of headbody length), and its fur is black and rather curly at the base, with a white tip. Adult males have a white patch of fur between their shoulders, red hair on back of their head, and white scrotum and nipples.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA5FFA0FFF36C27F5E2F89B.taxon	biology_ecology	Habitat. Steep mountainous areas in mixed deciduous and broadleaf evergreen or coniferous forest at elevations of 1500 - 2500 m. Temperatures here fall below freezing during five months of the year, and snow is common in winter. During this time, Guizhou Snubnosed Monkeys move down to rivers, as low as 570 m above sea level. They are sympatric with the Tibetan Macaque (Macaca thibetana) and the Rhesus Macaque (Macaca mulatta).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA5FFA0FFF36C27F5E2F89B.taxon	food_feeding	Food and Feeding. The diet of the Guizhou Snub-nosed Monkey contains mainly leaves (47 %), along with buds (15 %), flowers (9 %), fruits and seeds (22 %), and bark, bulbs, and some animal prey, mainly insect larvae (7 %). The diet varies by season. It contains mainly young leaves, flowers, unripe fruits, and seeds in spring, ripe fruits and seeds in summer, mature leaves in autumn, and buds in winter.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA5FFA0FFF36C27F5E2F89B.taxon	breeding	Breeding. Mating of the Guizhou Snub-nosed Monkey occurs between summer and autumn, with a peak in September. A single offspring is born in March — May. Infants are grayish-white and become darker gray as juveniles. Age at first birth is c. 9 years, with an interbirth interval of ¢. 3 years.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA5FFA0FFF36C27F5E2F89B.taxon	activity	Activity patterns. The Guizhou Snub-nosed Monkey is diurnal, arboreal, and terrestrial. Although primarily a tree-dweller, groups frequently cross open areas on the ground.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA5FFA0FFF36C27F5E2F89B.taxon	biology_ecology	Movements, Home range and Social organization. The Guizhou Snub-nosed Monkey lives in small unimale — multifemale groups of 3 - 10 individuals, or in bachelor groups of 2 - 5 individuals. In March — June and August-October, groups band together to form temporary conglomerations of over 400 individuals. Guizhou Snub-nosed Monkeys sleep huddled together in broadleaf evergreen trees, with dense foliage, to conserve heat. Their home ranges are ¢. 3500 ha.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA5FFA0FFF36C27F5E2F89B.taxon	conservation	Status and Conservation. CITES Appendix I. Classified as Endangered on The IUCN Red List. The Guizhou Snub-nosed Monkey is listed as Category I under the Chinese Wildlife Protection Act of 1989. It seems always to have been rare and was long known from only two specimens collected in 1903. In 2005, the total world population was estimated at 750 - 800 individuals. It is occasionally hunted (illegally) for its meat and use in traditional medicine, and it is sometimes caught in snares set for other animals. Habitat loss and disturbance are currently its major threats. The only place it is known to occur is Fanjingshan National Nature Reserve (China). Its extent of occurrenceis less than 5000 km? and the area of occupancyis less than 500 km?, in fewer than five locations. Tourism in this region is increasing considerably, with roads, hotels, and a cable car being constructed in the nature reserve. This and continued agricultural expansion and the collection of firewood have negative effects on the extant population. Due to its low genetic variability and restricted distribution, the Guizhou Snub-nosed Monkey might be highly vulnerable to environmental changes or climate oscillations. There is an urgent need to survey other remnant forests in the region, especially Jinfoshan National Nature Reserve, for the small possibility of finding other populations in the Wuling Mountains and also to investigate the possibilities for translocation. The most urgent conservation need is to remove threats in and around the Fanjingshan National Nature Reserve. Fanjingshan National Nature Reserve maintains a captive breeding colony, and a few pairs have been sent to other centers in China, but breeding is slow and the future of the captive population is not secure.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA5FFA1FAFF62BFF64BFCE0.taxon	materials_examined	China, Sichuan, near Moupin (= Baoxing) (30 ° 26 ’ N, 102 ° BOE).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA5FFA1FAFF62BFF64BFCE0.taxon	discussion	The subspecies of R. roxellana below differ from one another mainly in tail length, as well as in skeletal structure and width of the dental arch. Genetically, they do not \ :: segregate into reciprocal monophyletic clades. Three subspecies recognized.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA5FFA1FAFF62BFF64BFCE0.taxon	distribution	Subspecies and Distribution. R. r. roxellanaMilne-Edwards, 1870 — WCChina (SGanssu, SShaanxi, WSichuanprovinces). R. r. hubeiensisWang, Jiang & Li, 1998 — WCChina (ShennongjiainWHubeiProvinceandinNESichuanProvince). R. r. nlingensis Wang, Jiang & Li, 1998 — WC China (Qinling Mts, S Shaanxi Province).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA5FFA1FAFF62BFF64BFCE0.taxon	description	Descriptive notes. Head-body 56 - 83 cm (males) and 47 - 74 cm (females), tail 61 - 104 cm (males) and 51 - 92 cm (females); weight males 15 - 19 kg (occasionally up to 39 kg), females 6 - 10 kg (occasionally up to 20 kg). Coat of the Golden Snub-nosed Monkey is long and dense, being generally yellowish-red (varying from brown-red to bright golden-orange) and overlain with black on back. Limbs are similarly colored, with a thick black stripe running down outersides (although not to hands and feet) and a whitish patch on backs of thighs. Underside is yellowish-white, and crown is dark in adults. Males are similar to females but larger and more brightly toned, with a pronounced mantle, longer canine teeth, and a darker, more contrasting head and back. At the onset of winter, they shed summer coats and a thicker one begins growing. Penis is black, and scrotum is bluish-white. Both sexes of Golden Snub-nosed Monkey have violet-colored skin around their eyes and a broad, soft, whitish muzzle thatis sparsely haired; these two zones are separated, trefoil-like, by thickly furred wedges running from cheeks across bridge of the nose. Adult males develop swollen, reddish, fleshy flaps on corners of their upper lip, whose function (if any) remains a mystery. The “ Moupin Golden Snub-nosed Monkey ” (R. rn roxellana) is dull golden-red, with shoulders, forelimbs, and hindlimbs a dusky blackish-brown. Its tail is relatively short (i. e. about equal to head-body length), and its hindlimbs are relatively long compared with the other subspecies. In the “ Quinling Golden Snub-nosed Monkey ” (R. r. qinlingensis), pelage is uniformly brilliant golden-red. Its tail is relatively long (c. 115 % of head-body length). In the “ Hubei Golden Snub-nosed Monkey ” (R. » hubeiensis), dorsal pelage is somewhat paler (gray-brown) than on other subspecies, and its tail is considerably longer (i. e. ¢. 130 % of head-body length). In addition to pelage and morphological differences, subspecies differ in their nasal bones, premaxillae, and braincases; most strikingly, nasal bones, which are always short in this species, are absent on one or both sides in the Hubei Golden Snub-nosed Monkey.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA5FFA1FAFF62BFF64BFCE0.taxon	biology_ecology	Habitat. Montane conifer, deciduous and rhododendron forest, and bamboo thickets to elevations of 3000 m. The Golden Snub-nosed Monkeylives in a harsh environment where snow cover can last for up to six months. In summer, they inhabit regions above 3000 m, and in winter, they move down into valleys at 1800 - 2800 m. At Baihe, they occupy high-elevation conifer forests in winter. They occur in some of the same forests as the Giant Panda (Auluropoda melanoleuca). In the Qinling Mountains, the Quinling Golden Snub-nosed Monkey is found in mixed-deciduous broadleaf and mixed coniferous forest at 1400 - 2800 m. Moupin and Hubei golden snub-nosed monkeys occur most often in mixed coniferous and deciduous broadleaf forests.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA5FFA1FAFF62BFF64BFCE0.taxon	food_feeding	Food and Feeding. The Golden Snub-nosed Monkey is folivorous. It eats mainly lichens (29 %), fruits and seeds (29 %), and leaves (24 %), along with bark (11 %), buds, (4 %), twigs (1 %), and occasionally other items such as shoots, flowers, and animal prey (including insects, small vertebrates, bird eggs, and birds). Pine needles and pine cones are eaten during the leaner winter months. Diet composition varies seasonally; they feed more on mature leaves in summer, fruits and pine nuts in autumn, and lichens, bark, and buds in winter.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA5FFA1FAFF62BFF64BFCE0.taxon	breeding	Breeding. Female Golden Snub-nosed Monkeys have a 22 - 30 day reproductive cycle. Mating occurs in August — = October, with births in March — June, peaking in mid-April. A single offspring is born after a gestation of 194 - 203 days. Females usually give birth every two years. Newborns have long black hair on their head and back, and the eyes and nostril bridge are bluish-green. Infants are dark gray on their back and whitishgray below, with white hair on their ears. Their skin is white, except for the area around the mouth, which is pale pink, and palms and fingers are dark pink. Allomothering and infanticide have been observed. Adult males carry young on occasion. The age of sexual maturity for both sexes is 4 - 6 years. Interbirth intervals average 21 - 9 months. Life span in captivity is 25 - 30 years.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA5FFA1FAFF62BFF64BFCE0.taxon	activity	Activity patterns. Golden Snub-nosed Monkeys are diurnal and semi-arboreal. They spend a lot of time on the ground. The annual activity budget is 36 - 2 % resting, 35 - 8 % feeding, 22: 9 % moving, and 5 - 1 % engaging in other behaviors.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA5FFA1FAFF62BFF64BFCE0.taxon	biology_ecology	Movements, Home range and Social organization. The Golden Snub-nosed Monkey lives in unimale-multifemale or all-male groups, which vary in size according to season. In winter, smaller groups of 20 - 30 individuals are the norm, although they frequently come together into larger bands of up to 200 individuals in summer. Several bands may in turn combine temporarily to form enormous, fission-fusion troops of up to 600 individuals. Home ranges are 1000 - 4000 ha, but daily travel distance is generally less than ¢. 1000 m (shorter in winter than summer). The Golden Snub-nosed Monkey is adapted to the longest winter and coldest temperatures of any non-human primate. Although noted for their ability to tolerate extremely low temperatures, they nevertheless seem to have an aversion to snow. For this reason, they tend to migrate to low valleys at the onset of winter, during which time they also shed their summer coats and grow a new redder one. While traveling on the ground, bachelor males lead the group and bring up the rear. Natural predators include Dhole (Cuonalpinus), Gray Wolf (Canis lupus), Asian Golden Cat (Catopuma temminckii), Leopard (Panthera pardus), golden eagle (Aquila chrysaetos), and northern goshawk (Accipiter gentilis).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA5FFA1FAFF62BFF64BFCE0.taxon	conservation	Status and Conservation. CITES Appendix I. Classified as Endangered on The IUCN Red List, including all three subspecies. The Golden Snub-nosed Monkey is listed as Category I of the Chinese Wildlife Protection Act, 1989. It is by far the most common of the five species of snub-nosed monkeys. All three subspecies are threatened by habitat loss due to agriculture and tourism. The Moupin Golden Snub-nosed Monkey is also threatened by illegal hunting for meat, and sometimes hunters’ snares set for other animals. Due to increased protection, hunting has not been a major concern for Hubei and Quinling golden snub-nosed monkeys since 1990. Golden Snub-nosed Monkeys do not raid crops and are not, as such, an economic liability for people living near them. Development of tourism and the accompanying infrastructure such as roads has a significant negative impact on all three subspecies, most particularly Quinling Golden Snub-nosed Monkeys in the Qingling Mountains, and has already led to changes in the ranging behavior of groups in some areas. Fieldwork on the species has, until recently, typically used provisioning of favored foods to lure or even herd groups to observation sites; what effect this might have on their behavior is not known. Before 1990, they were threatened by illegal hunting, but this has stopped because of increased governmental protection. Golden Snub-nosed Monkeys have only rarely been displayed outside of China, and almost never outside of Asia, but they seem to do rather well in Chinese zoos, where they frequently breed. At one time, it was customary for Manchu mandarins and other high-ranking officials to wear robes made from woven fur of Golden Snub-nosed Monkeys. Aside from being a symbol of prestige, it was also thought to help ward off rheumatism. In 2006, the total population of the Moupin Golden Snub-nosed Monkey was estimated to be ¢. 10,000 individuals. A 2001 estimate put the total population of Quinling Golden Snub-nosed Monkeys at 3800 - 4000 individuals (c. 50 % of them mature), while a 1998 estimate for Hubei Golden Snub-nosed Monkeys was 600 - 1000 individuals with ¢. 250 or less mature individuals. Golden Snub-nosed Monkeys occur in at least eight protected areas in China: Moupin Golden Snub-nosed Monkey in Baihe and Wanglang nature reserves; Quinling Golden Snub-nosed Monkey in Changqing National Nature Reserve and Taibai, Zhouzhi, Foping, and Laoxiancheng nature reserves; and Hubei Golden Snub-nosed Monkey in Shennongjia Nature Reserve.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA4FFA1FA336675F94BF41F.taxon	materials_examined	26.43101 ° N, 98.38894 ° E (elevation 2815 m) in the Maw River area, northeastern Kachin State, in northeastern Myanmar.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA4FFA1FA336675F94BF41F.taxon	discussion	R. strykeri is closely related to R. bietu. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA4FFA1FA336675F94BF41F.taxon	distribution	Distribution. NE Myanmar (Salween-N’mai Hka divide in NE Kachin State, only around the Maw River, as far E as the Mts above the village of Chichitago, 26.31 ° - 26.51 ° N and 98.34 ° - 98.61 ° E) and S China (Gaolilgongshan National Nature Reserve, Yunnan Province).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA4FFA1FA336675F94BF41F.taxon	description	Descriptive notes. Head — body ¢. 55 - 5 cm (males) and c ¢. 54 cm (females), tail ¢. 78 cm (males) and ¢. 68 cm (females); weight c. 14 kg (males) and c. 8 - 5 kg (females). Fur of Stryker’s Snub-nosed Monkey is mostly black, with black or blackish ventral parts; limbs are all black or blackish. Crown has a thin, high, forward-curved crest of long, black hairs. There are protruding white ear tufts. Face is mostly naked and pale pink. Upperlips have whitish hairs on each side, and there is a distinct white beard. Perineal area is white. Tail is black and long (c. 140 % of head-body length). Stryker’s Snubnosed Monkey is most similar to the Yunnan Snub-nosed Monkey (R. bieti), from which it is distinguished by the total lack of white on underside and inner sides of limbs, lack of brown on temples, dark gray-brown forehead instead of buff, white whiskers instead of black, dark gray-brown hairs on cheeks instead of pure brown, shorter black hairs on backs of thighs instead of white, entirely pale pink facial skin, and a much longer tail (140 % of head-body length rather than 100 - 130 %).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA4FFA1FA336675F94BF41F.taxon	biology_ecology	Habitat. Steep mountainous areas in cool temperate rainforest and mixed temperate forests at elevations of 1720 - 3190 m and probably as high as 3660 m. Stryker’s Snub-nosed Monkey moves to high-elevation mixed temperate and conifer forests in summer (May — October) and to lower elevations in winter (November — April). It is sympatric with Shortridge’s Langur (Trachypithecus shortridger), various macaques (Macaca), and most probably the Bengal Slow Loris (Nycticebus bengalensis) and the Eastern Hoolock Gibbon (Hoolock leuconedys).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA4FFA1FA336675F94BF41F.taxon	food_feeding	Food and Feeding. Stryker’s Snub-nosed Monkeys eat bamboo shoots.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA4FFA1FA336675F94BF41F.taxon	breeding	Breeding. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA4FFA1FA336675F94BF41F.taxon	activity	Activity patterns. Stryker’s Snub-nosed Monkey is diurnal, arboreal, and terrestrial.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA4FFA1FA336675F94BF41F.taxon	biology_ecology	Movements, Home range and Social organization. Stryker’s Snub-nosed Monkeys have never been studied in the wild. Reports from the Maw River area indicated two groups of ¢. 30 individuals, one of 60 - 80, one of ¢. 80 - 100, and one of c. 150, but these may have been the same groups in different phases offission and fusion.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA4FFA1FA336675F94BF41F.taxon	conservation	Status and Conservation. CITES Appendix I. Classified as Critically Endangered on The IUCN Red List. The known population of Stryker’s Snub-nosed Monkey is estimated at 260 - 330 individuals. Its range is not more than ¢. 273 km? Illegal hunting is the major threat (at least 13 individuals were hunted and killed in 2009), and hunting and habitat loss will probably increase in the near future because of the construction of dams and access roads for logging. In 2011, Stryker’s Snub-nosed Monkey was discovered in Gaoligongshan National Nature Reserve near Pianma, Nujiang Lisu Prefecture, Yunnan, at an elevation of 2546 m in border forest contiguous with Myanmar. No population estimate is available for China.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA4FFA2FA3169E0F58BFABD.taxon	materials_examined	Indonesia, West Sumatra, South Pagai Island.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA4FFA2FA3169E0F58BFABD.taxon	discussion	The Pig-tailed Langur was originally described in its own monotypic genus, Simtas, although some later systematists chose to classify it in the genus Nasalis. Recent genetic data have suggested strongly that the closest relative of Simias is N. larvatus, the Proboscis Monkey of Borneo. Two subspecies recognized.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA4FFA2FA3169E0F58BFABD.taxon	distribution	Subspecies and Distribution. S. c. concolorG. S. Miller, 1903 — MentawaiIs (Sipora, NorthPagai, SouthPagaiIs, andnearbyIsofSimalegu, Simatapi, andSinakak); onlyprimateonSimaleguandSimatapi. S. c. stberu Chasen & Kloss, 1928 — Mentawai Is (Siberut I).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA4FFA2FA3169E0F58BFABD.taxon	description	Descriptive notes. Head — body 49 - 55 cm (males) and 46 - 52 cm (females), tail ¢. 16 cm (males) and c. 14 cm (females); weight 8.5 - 8.8 kg (males) and 7.7 - 2 kg (females). Male Pig-tailed Langursare larger and ¢. 30 % heavier than females, with canines ¢. 95 % longer in males than females. In both subspecies, there are two distinct color morphs. The first, more common, morph is dark grayish-black, with some agouti banding on anterior parts of the body, a white penal tuft, and a lighter-colored crown. The second morph is a pale buff color; it is somewhat uncommon on Siberut (c. 33 % of observed individuals in central and southern Siberut and only ¢. 5 % in northern Siberut) and extremely rare on the Pagai Islands. Both color morphs have white cheek patches and black on face, hands, and feet. The “ Pagai Pig-tailed Langur ” (S. c. concolor) is less dark, more brownish-toned, than the “ Siberut Pig-tailed Langur ” (S. ¢. siberu). White cheeks on the Siberut Pig-tailed Langur are more conspicuous than on the Pagai Pig-tailed Langur, and its black color (in the dark morph) has more banding on anterior parts. Fur on crown is directed backward, except for a pair of sideways-facing ear tufts. Nasal bone is elongated, but nose itself is snubbed and upturned. Tail is short, upturned, curly, and mostly hairless. The Pig-tailed Languris the only colobine with a short tail. Buttock pads of males are joined in the midline, but they are separated by the urogenital triangle in females. Unlike any other Asian colobine, female Pig-tailed Langurs exhibit sexual swellings, with the urogenital triangle swelling prominently during receptive periods of the menstrual cycle. Front and hind limbs are roughly equal length. Pig-tailed Langurs have a powerful odor.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA4FFA2FA3169E0F58BFABD.taxon	biology_ecology	Habitat. Swamp and lowland rainforest and primary hillside forest in the interior of the islands. Pig-tailed Langurs are sometimes found in mangrove forest but rarely in secondary forest. Mentawai forests are ever-wet with annual rainfall of up to 4000 mm.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA4FFA2FA3169E0F58BFABD.taxon	food_feeding	Food and Feeding. Pig-tailed Langurs eat mainly leaves, supplemented with fruits, seeds, and flowers. They occasionally raid cacao plantations.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA4FFA2FA3169E0F58BFABD.taxon	breeding	Breeding. Female Pig-tailed Langurs exhibit sexual swellings, but it is questionable whether they are comparable to swellings seen in most species of papionins and Chimpanzees (Pan troglodytes). Breeding appears to be seasonal, with mating in February-August and newborns most commonly seen in June-July. Single births are the norm.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA4FFA2FA3169E0F58BFABD.taxon	activity	Activity patterns. The Pig-tailed Langur is diurnal, arboreal, and terrestrial. When alarmed by humans, they remain motionless in dense foliage, not making a sound. If threatened further, they drop to the ground and attempt to flee.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA4FFA2FA3169E0F58BFABD.taxon	biology_ecology	Movements, Home range and ‘ Social organization. Despite the fact that the limbs of Pig-tailed Langurs are roughly of equal length (suggesting a more terrestrial existence), their movements are mostly through trees. Group travel is initiated by adults of either sex, with no consistent leader, and group members stay within a short distance of each other throughout the day. They tend to move more slowly and quietly than sympatric Presbytis. Males produce loud calls (a series of loud nasal barks), mostly in the early morning but also throughout the day, with another peak in the late afternoon. Females with infants often “ park ” them in a tree crown while feeding nearby. Groups of Pig-tailed Langurs typically sleep in densely foliated trees in the canopy or in emergent trees. Home range size is variable, depending on their density. In most areas, typical home ranges are c. 7 - 25 ha. Hunting appears to affect group size and social organization by changing densities. Social organization is variable, described as “ monandrous, ” because the most commonly observed group composition is one adult male and one or more adult females. Multimale — multifemale groups have also been reported, with more adult females than males. Average group size for the Siberut Pig-tailed Langur in northern Siberut (7 - 9 individuals) is reportedly higher than in other parts of Siberut or for the Pagai Pig-tailed Langur throughout its range (3 - 4 individuals). Population densities of the Pagai Pig-tailed Langur are ¢. 5 - 21 ind / km? in unlogged forest. The density of the Siberut Pig-tailed Langur is up to 53 ind / km? in the Peleonan Forest in northern Siberut. The Pig-tailed Langur is sympatric with the Siberut Langur (Presbytis siberu), the Siberut Macaque (Macaca siberu), and Kloss’s Gibbon (Hylobates klossii) on Siberut and with the Mentawai Langur (FP. potenziani), the Pagai Macaque (Macaca pagensis), and Kloss’s Gibbon on the southern Mentawai Islands. Pig-tailed Langurs form mixed-species associations with the Mentawai Langur and the Siberut Langur.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA4FFA2FA3169E0F58BFABD.taxon	conservation	Status and Conservation. CITES Appendix I. Classified as Critically Endangered on The IUCN Red List, including both subspecies. The Pig-tailed Langur is protected by Indonesian law. They are remarkably little known or studied; indeed, up until the late 1970 s, they had not even been photographed. They are threatened mainly by hunting for meat, commercial logging, conversion of their habitat into oil palm plantations, and forest clearing by local people. While all Mentawai primates are hunted, Pig-tailed Langurs are preferred prey in all areas. Recently, hunting has increased because of improved access to remote areas due to logging roads and tracks, replacement of bows and arrows with rifles, and changes in local rituals and taboos that formerly regulated hunting. Pig-tailed Langurs are sometimes taken for the pet trade, but they rarely, if ever, thrive. They are also occasionally poisoned as agricultural pests. All of these factors have resulted in a significant decline in their numbers and even their total elimination on some islands that they formerly occupied. The Siberut Pig-tailed Langur has the largest remaining population of 6000 - 15,500 individuals, estimated in 2006. The Pagai Pig-tailed Languris in urgent need of protective measures, with a total population estimated at just 700 - 1800 in 2006. The most recent estimate of total population size for the Pig-tailed Langur was 6700 - 17,300 in 2006, representing a decline of 73 - 90 % since a 1994 estimate. It is known to occur in only one protected area (Siberut National Park), butit also occurs in the proposed Betumonga Research Area on North Pagai (recently reported to have been logged over). It is found at its highest density in the Peleonan Forest in northern Siberut, which, as of 2011, is protected on a short-term basis by local agreements to exclude logging concessions. In 2006, the following conservation actions were suggested: increased protection for Siberut National Park, which currently lacks enforcement; formal protection of the Peleonan Forest in North Siberut, which has an unusually large primate population and is easily accessible; protection of areas in the Pagai Islands by cooperating with a logging corporation that has practiced sustainable logging techniques since 1971; conservation education, especially regarding hunting; and the development of alternative economic models for local people to reduce the likelihood of them selling off their lands to logging companies.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA7FFA3FAEF635FF809F871.taxon	materials_examined	Pontiana (= Pontianak), Borneo.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA7FFA3FAEF635FF809F871.taxon	discussion	Two subspecies of N. larvatus are sometimes recognized: larvatus and orientalis. There is little difference between them, however, and the latter is not recognized by all authorities. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA7FFA3FAEF635FF809F871.taxon	distribution	Distribution. Borneo (Sabah and Sarawak states, Brunei, and Kalimantan); it is also found on the satellite islands of Berhala, Sebatik, and Pulau Laut.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA7FFA3FAEF635FF809F871.taxon	description	Descriptive notes. Head — body 73 - 76 cm (males) and 61 - 64 cm (females), tail 66 67 cm (males) and 55 - 62 cm (females); weight 20 - 24 kg (males) and c. 10 kg (females). The Proboscis Monkey is strongly sexually dimorphic. It is a highly distinctive large, predominantly chestnut-brown monkey, with bare pink to brown faces. Ventral fur is paler than dorsal fur, except around sides of face and an almost complete yellowish collar around neck. Lower limbs, hands, and feet are gray. Stomach is relatively twice as large as any other species of colobine, so its abdomen appears bloated. The male is especially striking with his uniquely enormous, pendulous nose, dark chestnut cap, thick darker brown fur across upper and middle back and sides, and white rump patch and tail. The female has a smaller, “ snubbier ” nose, and her body is more uniformly brown, but with a gray rump patch and tail. Neonates have sparse, blackish fur and blue faces, with snubby upturned noses; by c. 4 months of age, fur has turned brown, but face retains a blue-gray tinge for up to a year. Proboscis Monkeys have long hind feet, with partially webbed digits.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA7FFA3FAEF635FF809F871.taxon	biology_ecology	Habitat. Almost exclusively in coastal forests and forests close to major rivers, especially mangrove, peat swamp, and riparian forests. Proboscis Monkeys occur farther inland immediately adjacent to large rivers in Kalimantan and in inland swamps of Danau Senatarum and Mahakam Lakes. It is never found above elevations of 350 m, except for extremely rare seemingly vagrant individuals. It also occurs in some lowland dipterocarp forests but only near to rivers. In all habitats, Proboscis Monkeys are rarely found more than c. 1 km from a significant waterway. Formerly, they were widespread throughout most coastal forests of Borneo and possibly along most major rivers.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA7FFA3FAEF635FF809F871.taxon	food_feeding	Food and Feeding. Studies of Proboscis Monkeys at three sites in different parts of Borneo concur that the diet predominantly comprises young leaves (38 - 66 %) and unripe fruits and seeds (26 - 50 %), with flowers only ¢. 3 %. They occasionally forage on tree bark and termite nests. In Kinabatangan, Sabah, Proboscis Monkeys feed on 188 different plant species. Diets vary seasonally in food item type and diversity, depending largely on availabilities of edible fruits and seeds. In Samunsam, Sarawak, when fruits in riparian forests are scarce, they feed predominantly on young leaves and shoots of Rhizophora (Rhizophoraceae) species in mangroves.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA7FFA3FAEF635FF809F871.taxon	breeding	Breeding. Proboscis Monkeys give birth to single offspring after a gestation of ¢. 166 days. Sexual maturity is reached at 3 - 5 years in females and 5 - 7 years in males. Individual females generally breed every other year. In the wild, births can occur throughout the year but with a pronounced peak during the rainy season in November — February. Infants andjuveniles commonly scream and display at mating adults, including pulling on a mating male’s leg or nose.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA7FFA3FAEF635FF809F871.taxon	activity	Activity patterns. The Proboscis Monkey is diurnal, arboreal, and sometimes terrestrial. In almost all parts of their distribution, they spend the night in trees immediately adjacent to significant waterways, moving up to 750 m inland during the day; they return by late afternoon to a river prior to sleeping there for the night. As different groups move toward and assemble beside a river in the evenings, adult males give frequent spectacular displays, leaping between trees often with loud roars and landing with a noisy crash. One exception is at Bako National Park, Sarawak, where individuals spend the night in trees on cliffs slightly inland, moving down to mangrove bays in the early morning. Activity throughout the day largely comprises feeding periods interspersed with prolonged resting / digesting periods.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA7FFA3FAEF635FF809F871.taxon	biology_ecology	Movements, Home range and Social organization. Daily movements of Proboscis Monkeys are determined strongly by the distribution of waterways, and their sleeping sites are generally beside open water. Predators include crocodiles (Crocodylus), Diard’s Clouded Leopards (Neofelis diardi), and reticulated pythons (Python reticulatus). Where waterways are narrow, Proboscis Monkeys traverse them by arboreal routes. They are also proficient swimmers. For widerrivers, they leap from a high branch into the river and then swim the final stretch or, for the wider rivers, climb down to the bank and slip into the water to swim across, with infants and females generally in front and the male farther behind. Proboscis Monkeys can swim considerable distances under water if disturbed. Data on daily movement patterns of individual groups are scarce, but they generally travel less than 1000 m. Home range sizes vary between sites, depending on habitat type and distribution of waterways. In heterogeneous lowland and mangrove forests of Samunsam, Sarawak, home ranges are c. 9 km? whereas in the peat swamp forests Tanjung Puting, Kalimantan, they are 1 - 4 km ®, in both cases encompassing forest on both sides of the river. Proboscis Monkey groups are non-territorial, with more than 90 % home range overlap. Social groups comprise a single adult male and up to nine adult females and their offspring. Ranges of these unimale — multifemale groups overlap, and they frequently aggregate, especially beside rivers in the evenings. Both males and females transfer between groups. Males leave their natal groups at c. 2 years of age and generallyjoin all-male groups, which in most cases are of similar size as unimale — multifemale groups. Infanticide has never been recorded, but occasional movements of females with dependent infants out of their unimale-multifemale groups indicate that it might occur.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA7FFA3FAEF635FF809F871.taxon	conservation	Status and Conservation. CITES Appendix I. Classified as Endangered on The IUCN Red List. Proboscis Monkeys are fully protected by law in all of the Bornean states where they occur, although enforcement is often weak. Coastal lowlands and land adjoining large rivers, which are the original habitats of the Proboscis Monkey, are the most developed areas of Borneo: most people live along rivers, most large towns are near river mouths, and most large-scale agriculture and aquaculture schemes are on coastal and alluvial plains. Few large, intact areas of suitable habitat remain, and populations of Proboscis Monkey are now highly fragmented and scattered; in Sabah, only an estimated 9 - 8 % of the original suitable habitat remains. Of the remaining habitat, mangroves in many areas have been subjected to intensive logging for poles and charcoal. Peat swamp forests have also been heavily and repeatedly logged in many areas. In the 1970 s and 1980 s, such logging was sometimes followed by poisoning of non-timber trees, many of which were food trees for wildlife, with the aim of enhancing the next timber crop. This practice has been discontinued, but it was responsible for major degradation of some peat swamp forests. Forest fires along rivers can be damaging; e. g. the 1997 - 1998 Bornean fires destroyed large areas of remaining habitat. Hunting in the past might have been significant in some areas, potentially accounting for the limited and patchy distribution of the Proboscis Monkey along major rivers. In coastal areas, traditionally, hunting was not a significant problem because inhabitants are predominantly Muslims who do not eat monkeys. With the advent of speedboats and shotguns, however, people now come from towns to hunt Proboscis Monkeys for sport; their habit of sleeping conspicuously next to rivers makes them easy targets. Intense pressure on coastal land means that few reserves are large enough to protect whole populations, and the current status of the Proboscis Monkey is not well known. It is thought to occur in 18 protected areas, although all but five are too small, or habitat too marginal, to fully protect their populations. The protected areas are: Samunsam Reserve, Bako National Park, Kuching Wetland National Park, Sedilu National Park, Maludam National Park, and Rajang Mangrove National Park in Sarawak, Malaysia; Pulau Siarau & Pulau Beramban Primary Conservation Areas and Bukau Api Api Protection Forest Reserve in Brunei Darussalam; Maliau Basin Conservation Area, Danum Valley Conservation Area, Sepilok Orang Utan Sanctuary, Lower Kinabatangan Wildlife Reserve, and Kulamba Wildlife Reserve in Sabah, Malaysia; and Kutai National Park, Pleihari Tana Laut Wildlife Reserve, Tanjung Puting National Park, Gunung Palung National Park, and Danau Sentaram National Park in Kalimantan, Indonesia. Recent estimates indicate that 6000 Proboscis Monkeys remain in Sabah, mostly in disjunct populations in the Kinabatangan and Segama river floodplains, with a smaller, fragmented population on the Klias Peninsula on the west coast. In Sarawak, less than 1000 individuals remain in patchily distributed populations. Proboscis Monkeys are more abundant in Kalimantan, with population sizes of 100 - 1000 individuals, although recent surveys are lacking. The population in the Mahakam Delta, which would have numbered in the thousands until the early 1990 s, has been decimated because of conversion of the coastal swamps to shrimp farms. The Proboscis Monkey is now extinct in Indonesia’s Pulau Kaget Nature Reserve where it was once abundant.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA8FFAEFF3664A3FD6CFD7C.taxon	materials_examined	India, Bengal.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA8FFAEFF3664A3FD6CFD7C.taxon	discussion	S. entellus shows considerable variation in size and coat color throughout its distribution. For many years, the Hanuman or gray langurs of South Asia were all classified as subspecies of S. entellus, but W. C. O. Hill in his 1939 review and C. P. Groves in his 2001 Primate Taxonomy considered it to be monotypic, and this has been confirmed with recent genetic studies. M. L. Roonwal separated gray langurs of South Asia into a northern group and a southern group based on tail carriage. S. entellus is of the northern group (Type IA), with the tail looping forward and dangling down and the tip reaching back, even hanging down on one side of the body. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA8FFAEFF3664A3FD6CFD7C.taxon	distribution	Distribution. India, the species’ range extends 1900 km W to E, and includes all populations with the northern-type tail carriage between the Tapti River in Gujarat State and Krishna River in Andhra Pradesh State to the foothills of the Himalaya. Introduced into SW Bangladesh, where the population may have arisen from descendents of a pair introduced by Hindu pilgrims on the banks of the Jalangi River.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA8FFAEFF3664A3FD6CFD7C.taxon	description	Descriptive notes. Head — body 45.1 - 78.4 cm, tail 80.3 - 111.8 cm; weight 16.9 - 19.5 kg (males) and 9.5 - 16.1 kg (females). Coat of the Bengal Sacred Langur has short, wavy hairs that are creamy-yellow on head and flanks and brown on back and limbs. Tail has a white tip. Underside is red-gold. Cheek whiskers are long and whitish. Face and ears are black. Eyebrows are long and bristle-like. Crown has no tuft, and hairs are parted neatly along midline. Hands and feet are black, in contrast to brownish limbs.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA8FFAEFF3664A3FD6CFD7C.taxon	biology_ecology	Habitat. Mostly dry tropical deciduous forest, tropical thorn forest in parts of Gujarat and Rajasthan; tropical moist deciduous forest on the eastern and south-eastern edges of its distribution; and human-modified sparsely wooded areas and scrublands. Bengal Sacred Langurs are often found in proximity to human habitation. They are a lowland plains species; their ranges extend up to elevations of 400 m.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA8FFAEFF3664A3FD6CFD7C.taxon	food_feeding	Food and Feeding. Bengal Sacred Langurs feed on leaves, shoots, fruits, buds, flowers, tree exudates, bark, vines, grasses, galls and insect larvae, and termite soil. At Kanha, Madhya Pradesh, the annual diet of a group (time spent feeding on different food items) was 39 % leaves (mostly mature 35 %; young leaves 4 %), 24 % seeds and fruits, 9 % flowers, 11 % buds and bark, and 3 % other items including insects. Caterpillars and other insects can make up to 25 % of their diet in monsoon months. No single species predominated throughout the year. The group exploited 53 species of trees and vines. Young leaves were eaten in preference to mature leaves, and fruits and flowers in preference to leaves. The ability of Bengal Sacred Langurs to survive on diets of largely mature leaves is fundamental to their survival in dry seasons when canopies of deciduous forests become leafless and only understories remain green. They play an important role in seed dispersal of trees such as Aglaia lawii (Meliaceae). They feed on bark of Boswellia serrata (Burseraceae), Ficus glomerata (Moraceae), Anogeissus latifolia (Combretaceae), Albizia odoratissima (Fabaceae), and Tamarindus indica (Fabaceae), especially during dry seasons when food is limited. They also feed on cultivated crops.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA8FFAEFF3664A3FD6CFD7C.taxon	breeding	Breeding. Breeding and mating of the Bengal Sacred Langur occur throughout the year when food is abundant, and especially when they are provisioned in temples, urban areas, or villages, or have access to crops. In forest groups and more seasonal areas, the birth season is in December — May. Seasonality in conceptions is sometimes apparent, but this is an effect of varying food availability. Only females in sufficient physical condition conceive. Males and females reach puberty at c. 4 years of age but are not fully developed until 6 - 7 years. The menstrual cycle is 25 - 27 days. Females show proceptive behavior; shaking their heads, hopping, lowering their tails, and presenting their hindquarters in front of males. The gestation period is ¢. 200 days (189 - 208 days), and the interbirth interval is ¢. 16 - 7 months. At Jodhpur, Rajasthan, females have their first infant when they are c. 3 - 5 — 4 years old. Infants are usually born single. A mother suckles her infant for 12 - 13 months. Females other than the mother carry and care for infants (alloparenting) for their first month or so. From birth to ¢. 5 months, infants are brown, but from then to c. 12 - 15 months, their pelage changes to a pale off-white and then to that of adults. Males are considered adult at 6 - 7 years old. Bisexual groups atJodhpur and Abu are not closed in terms of breeding. All-male groups are frequently near mixed groups and often able to associate closely with receptive / proceptive females. While rapid all-male replacement and subsequent infanticide (to make females return to ovulatory cycling and sexual receptivity) by groups of males happens at Abu and Jodphur, male replacement also occurs as a gradual process over 2 - 3 months, without takeovers and infanticide. Temporary associations of non-group adult males and females in bisexual groups are more common than changes in male membership involving actual replacement. As is typical of colobines, resident males are passively tolerant of infants but defend them when they are threatened. At Orcha, Madhyar Pradesh in the south-western part of their distribution, male Bengal Sacred Langurs are indifferent to male infants until they are ¢. 10 months old. A male infant at this age will run up to a passing male, mountit, and then, as its dismounts, the male sits and the infant runs round to face him and they embrace. This behavior is displayed by young males until they are c. 4 years old.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA8FFAEFF3664A3FD6CFD7C.taxon	activity	Activity patterns. Bengal Sacred Langurs are diurnal and primarily terrestrial. The activity budget of a group at the Kanha Tiger Reserve, Madhya Pradesh, was 41 - 8 % inactive (resting), 25 - 7 % feeding, 13 - 1 % traveling, and 19 - 4 % engaging in social and other activities. When fleeing, they tend to run along the ground rather than through trees. During a total eclipse, Bengal Sacred Langurs go into a state of total dormancy and inactivity, regaining normality only when the eclipse is over.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA8FFAEFF3664A3FD6CFD7C.taxon	biology_ecology	Movements, Home range and Social organization. Bengal Sacred Langurs live in one-male or multimale bisexual groups and all-male groups. Group sizes are usually 30 - 80 individuals. At the Sariska Wildlife Refuge in Rajasthan, they are 30 - 125 (average 64), and the male-to-female sex ratio is 1: 2 - 6. At the Kanha Tiger Reserve, unimale — multifemale groups can be as large as 34 individuals, with up to 15 adult females. Most males there live in unimale — multifemale groups, with an average of 22 individuals. In Jodhpur, all males are either solitary, live in unimale-multifemale, or all-male groups; average size is 38 individuals, with an adult sex ratio of 1: 4 - 9. At Orcha, maleslive in either multimale-multifemale or all-male groups, but not in unimale-multifemale groups; average size is 19 individuals, with an adult sex ratio 1: 1 - 6. Large groups are associated with more open habitats, their degree of terrestriality, and increased threats from ground predators such as Leopards (Panthera pardus) or Tigers (P. tugris). When not feeding, males, not females, tend to sit up in trees, vigilant for other langur groups and predators. Related to this, Chital (Axis axis) are often found in association with Bengal Sacred Langurs. Chital exploit the vigilance of male langurs, while langurs undoubtedly benefit from the enhanced sense of smell and hearing in the Chital. They respond to each others’ alarm calls. In these large groups, males show no strict dominance hierarchy and are quite tolerant of each other. Females are philopatric, they stay in the group they were born in, but males disperse and become temporarily solitary or join up with all-male groups. Solitary males are healthier than males living in groups. Females groom males and other females, but males very rarely groom females. Females have a strict dominance hierarchy among themselves that is influenced by age. Younger females are higher ranking, and they give and receive more grooming than older females and those lower in the hierarchy. Females in conflict show a curious behavior ofsitting erect, face-to-face, while slapping each other with both hands. Male group sizes are 2 - 43 individuals, with a mean of 21. Home ranges are 40 - 90 ha. Home range sizes are influenced by temporal and spatial variation in abundance and dispersion of food sources but also by proximity to agricultural crops, food provisioning, and other environmental factors. Daily movement can be more than 1000 m. Bengal Sacred Langurs travel further when their diets are more focused on fruits and flowers than leaves. At least in some populations (Abu, Jodhpur, and Kanha), male takeovers (invasion of bisexual groups by nongroup males, eviction of resident group males, and annexation of the group’s females and immature members) are sometimes rapid and complete and associated with subsequent infanticide. In other populations (Kaukori, Sariska Wildlife Sanctuary, and Gir Wildlife Sanctuary), male membership of groups changes gradually, without involving abrupt takeovers and infanticide. At Kanha Tiger Reserve, male tenure as a dominant in a group averaged c. 45 months. Males in unimale — multifemale groups at Jodhpur manage an average tenure of 2 - 2 years. Adult males may leave the group and live alone or join all-male groups. Bengal Sacred Langurs are skilled at raiding crops, with strategic raiding operations where a few individuals act as guards and the rest of the group feeds. Robust male members of unimale and multimale bisexual groups and all members of all-male groups are more frequent crop raiders than females. Females with infants enter fields to raid crops only when males are with them to act as guards. Densities of Bengal Sacred Langurs vary considerably in different populations. At Orcha, Madhya Pradesh, in the south-western part of their distribution, they can be as low as 5 ind / km?, but they reach 46 ind / km? in Kanha Tiger Reserve, north of Orcha, and 121 ind / km? in Gir Forest.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFA8FFAEFF3664A3FD6CFD7C.taxon	conservation	Status and Conservation. CITES Appendix I. Classified as Least Concern on The [UCN Red List. The Bengal Sacred Languris listed on Schedule II, Part I of the Indian Wildlife (Protection) Act, 1972, amended up to 2002, and Schedule III, Bangladesh Wild Life (Preservation) Order, 1973. The Bengal Sacred Langur occurs in many protected areas. Its status in captivity is difficult to determine because of taxonomic confusion with related forms. It is listed as Least Concern in view of its wide distribution, tolerance of a broad range of habitats, and presumed large population. If threats arising from hunting and human-animal conflicts increase (e. g. in Andhra Pradesh and Orissa), the Bengal Sacred Langur will need to be reassessed and could qualify for listing as Near Threatened. Its total population size is unknown. Most Bengal Sacred Langurs now occupy human-dominated landscapes, with very few actually occurring in forested areas. Habitat loss, intensive agriculture, forest fires, and conflict with humans (including road kills) are major threats. They like to stay and move close to roads or on them because roads are avoided by predators such as feral dogs and leopards. Bengal Sacred Langurs are revered and fed in temples.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFABFFAEFFE067D8FC6FF52C.taxon	materials_examined	India, Chamba, Deolah, 1800 m above sea level.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFABFFAEFFE067D8FC6FF52C.taxon	discussion	S. ajax was listed by W. C. O. Hill in his 1939 review of Asian langurs as a subspecies of S. schistaceus. C. P. Groves in his 2001 Primate Taxonomy considered ajax sufficiently distinct as to warrant the status of species. Genetic analyses may yet indicate a revision of this arrangement. M. L. Roonwal separated gray langurs of South Asia into a northern group and a southern group based on tail carriage. S. ajax is of the northern group (Type IA), with the tail held looping forward and dangling down, even hanging down on one side of the body. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFABFFAEFFE067D8FC6FF52C.taxon	distribution	Distribution. NW India, confined to the Chamba Valley of Himachal Pradesh State; it may also occur in the Kishtwar Valley ofJammu and Kashmir State.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFABFFAEFFE067D8FC6FF52C.taxon	description	Descriptive notes. Head-body 50.8 - 78.7 cm, tail 72: 4 - 96.5 cm; weight 19.5 - 20.8 kg (males). The Chamba Sacred Langur has yellowish upper sides and yellow-tinged under sides that characterize its loose shaggy coat. Head is white and is less differentiated from rest of body, as in other congeners. Tail tip is white. Arms are dark below elbows, and patches of dark skin are evident on hands. Legs and body are similarly colored, and feet are a little darker. Face and ears are black, and eyebrows are long and bristle like. Crown is neatly parted along the midline.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFABFFAEFFE067D8FC6FF52C.taxon	biology_ecology	Habitat. Himalayan moist temperate, semi-evergreen, and subalpine cedar forest and alpine scrub at elevations of 2200 - 4000 m. Chamba Sacred Langurs can also be seen near villages.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFABFFAEFFE067D8FC6FF52C.taxon	food_feeding	Food and Feeding. There is no specific information available for this species, but the Chamba Sacred Languris presumably similar in its diet and feeding habits to the other Himalayan species such the Nepal Sacred Langur (S. schistaceus) and the Terai Sacred Langur (S. hector).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFABFFAEFFE067D8FC6FF52C.taxon	breeding	Breeding. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFABFFAEFFE067D8FC6FF52C.taxon	activity	Activity patterns. The Chamba Sacred Langur is diurnal and mainly terrestrial.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFABFFAEFFE067D8FC6FF52C.taxon	biology_ecology	Movements, Home range and Social organization. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFABFFAEFFE067D8FC6FF52C.taxon	conservation	Status and Conservation. CITES Appendix I. Classified as Endangered on The IUCN Red List. The Chamba Sacred Languris listed on Schedule II, Part I of the Indian Wildlife (Protection) Act, 1972, amended up to 2002. Occurring in a single restricted locality, it 1 s also recognized under the Alliance for Zero Extinction (AZE), as an AZE Species, attracting specific and urgent measures forits conservation. Population estimates indicate only 250 mature adults and less than 500 individuals confined to the narrow Chamba Valley in Himachal Pradesh. Some of the imminent threats confronting the Chamba Sacred Langur include habitat loss, retaliatory killing due to crop raiding, persecution, hunting, logging, expansion of commercial horticulture, and change in land use and other development activities.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFABFFAEFFE06E04F6C7F32C.taxon	materials_examined	India, Sitabani, Ramnagar, Kumaon, 600 m above sea level.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFABFFAEFFE06E04F6C7F32C.taxon	discussion	S. hector was listed by W. C. O. Hill in his 1939 review of Asian langurs as a subspecies of S. schistaceus. C. P. Groves in his 2001 Primate Taxonomy considered hector sufficiently distinct as to warrant the status of species. Genetic analyses may yet indicate a revision of this arrangement. M. L. Roonwal separated gray langurs of South Asia into a northern group and a southern group based on tail carriage. S. hector is of the northern group (Type IB), with the tail looping forward but held up above the back with the tip out-curved. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFABFFAEFFE06E04F6C7F32C.taxon	distribution	Distribution. Himalayan foothills in N India (states of Uttarakhand, Uttar Pradesh, and W Bengal), and W & C Nepal; it also probably occurs in SW Bhutan near Pankhabari.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFABFFAEFFE06E04F6C7F32C.taxon	description	Descriptive notes. Head — body 58.4 - 76.2 cm, tail 75 - 99.1 cm; weight 17 kg (males) and 17 - 7 kg (females); these measurements were taken from a sample in which the Terai Sacred Langur and the Nepal Sacred Langur (S. schistaceus) were considered synonymous. Coat of the Terai Sacred Langur is long and thick, with a plush grayish-yellow to pale orange over body and a dark line running down back. Underside is bufty-white. Fur acquires a brownish tinge below elbows and knees. Hands and feet are pale. Head is a striking white, and cheek whiskers are long and whitish, forming a bushy halo around the jet-black face. Crown is parted neatly along the midline. Tail tip is white.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFABFFAEFFE06E04F6C7F32C.taxon	biology_ecology	Habitat. Semi-evergreen sal (Shorea robusta, Dipterocarpaceae) forest, moist deciduous forests of the Siwalik Hills, and alpine cedar and mid-elevation oak forest in the Himalayan foothills, and the Terai (a belt of marshy grasslands savannas and forests south of outer foothills of the Himalaya and Siwalik Hills, and north of the Indo-Gangetic Plain, at elevations of 150 - 1600 m). Winter weather is harsh, with snowstorms and temperatures below freezing in December — February.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFABFFAEFFE06E04F6C7F32C.taxon	food_feeding	Food and Feeding. Terai Sacred Langurs eat flowers and flower buds, unripe and ripe fruit, seeds, buds, young and mature leaves, petioles, pith, bark, wood, gum, insects (including their excreta), and soil. In year-long studies by A. Koenig and C. Borries and colleagues at Ramnagar, 64 % of feeding time was spent on leaves, 15 % fruits, 6 % flowers, 5 % other plant parts, and 10 % on insects and soil. Sixty-nine plant species contributed 90 % of the diet, and of these, 57 contributed less than 1 % to the diet, a further nine contributed 19 - 2 %, and just three species contributed 54 %. Particularly important were Spatholobus parviflorus (Fabacae), a leguminous vine that provided flowers, fruits and leaves, Terminalia bellirica (Combretaceae), and Dillenia pentagyna (Dillenicaeae). Terai Sacred Langurs also feed on crops and in orchards.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFABFFAEFFE06E04F6C7F32C.taxon	breeding	Breeding. The Terai Sacred Langur is a seasonal breeder; births occur only in January — June, with nearly 50 % of all births occurring in March. Mating is frequent in May-June, but conceptions are restricted to July-November, with ¢. 50 % occurring in August. The gestation period is ¢. 209 days. The menstrual cycle averages 26 - 4 days. Females give no visible sign of menstruation. Females are proceptive; they present their anogenital region to males, shaking their head and lowering their tail. The mean age atfirst birth is 6 - 7 years (range 6 - 8). Lactation lasts ¢. 25 months, but it can be as long 31 months. Sixty-five percent of females weaned their infants only when they were pregnant again. Studies at Ramnagar have shown that males can, infrequently, be infanticidal after immigrating into a new group. When an infant is raised successfully to independence, the average interbirth interval is 2 - 4 years. The intervalis halved (1 - 2 years) if the infant dies before it is one year old, and reduced to two years if it dies in its second year. In five groups at Ramnagar, male infanticide accounted for 30 - 8 - 62 - 5 % of infant mortalities during the first two years of life. Females copulate even when they are pregnant, supposedly so as to confuse paternity and protect their offspring from infanticide. Nevertheless, most attackers (91 %) were resident males and killed infants despite having copulated with the mother if they had not been a member of the group when the infant was conceived or born.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFABFFAEFFE06E04F6C7F32C.taxon	activity	Activity patterns. Terai Sacred Langurs are diurnal, arboreal, and terrestrial. At Ramnagar, they spend ¢. 20 % of their day on the ground and the remainder in trees.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFABFFAEFFE06E04F6C7F32C.taxon	biology_ecology	Movements, Home range and Social organization. Groups of Terai Sacred Langurs are unimale — multifemale or (mostly) multimale-multifemale. Group sizes average 18 - 3 individuals (range 7 - 41), with an average of 2 - 5 adult males and 6 - 6 adult females. About 72 % of bisexual groups have more than one adult male (as many as nine). Females are philopatric, they remain in their natal groups. Males disperse at c. 6 years old when they are still considered subadults. Females form a linear and stable dominance hierarchy, with younger adult females being the most dominant; older females gradually drop down in the hierarchy. Males do not form permanent all-male bands as is found in other gray langurs. They remain solitary, form pairs, or form ephemeral all-male groups of up to five individuals. Male replacement involves staggered immigrations and exclusions, and it is gradual. A dominant male’s tenure can last six or more years. Male Terai Sacred Langurs that are fathers or were resident when infants are born are highly protective of them. When entering a new group, they do it on their own and establish their dominance within a few days. The resident male either stays in the group as a subordinate or is expelled. The density of Terai Sacred Langurs at Ramnagar has been estimated at 26 ind / km?. In the Khumaon Hills, Himalaya, the density is 97 ind / km?.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFABFFAEFFE06E04F6C7F32C.taxon	conservation	Status and Conservation. CITES Appendix I. Classified as Near Threatened on The IUCN Red List. The Terai Sacred Langur is listed under the Nepal National Park and Wildlife Conservation Act (1973) as a common animal. It occurs in three protected areas: Corbet Tiger Reserve, Rajaji National Park, and Valmiki National Park in India and several protected areas in Nepal. The population is threatened by development activities such as mining and firewood collection, charcoal collection and production, habitat loss due to human settlements, and electrocution from powerlines.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFABFFAFFAEA6809F9F4F9F5.taxon	materials_examined	Nepal.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFABFFAFFAEA6809F9F4F9F5.taxon	discussion	In his 1939 review of the Asian langurs, W. C. O. Hill placed the Himalayan forms hector, achilles, lanius, and ajax as subspecies of S. schistaceus. C. P. Groves in his 2001 Primate Taxonomy listed lanius and achalles as junior synonyms of S. schistaceus, but he considered hector and ajax sufficiently distinct as to warrant the status of species. Genetic analyses may yet indicate a revision of this arrangement. M. L. Roonwal separated gray langurs of South Asia into a northern group and a southern group based on tail carriage. S. schistaceusis of the northern group (Type IB), with the tail looping forward but held up above the back with the tip out-curved. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFABFFAFFAEA6809F9F4F9F5.taxon	distribution	Distribution. NW Pakistan, N India (Jammu and Kashmir, Himachal Pradesh, Uttarakhand, and NW Bengal states, and Sikkim), S China (Tibetan regions of Bo Qu, Ji Long Zang Bu and Chumbi Valleys in Xizang Autonomous Region), Nepal, and W Bhutan (E to Sankosh River); its presence in E Afghanistan is uncertain.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFABFFAFFAEA6809F9F4F9F5.taxon	description	Descriptive notes. Head-body 58.4 - 76.2 cm, tail 75 - 99.1 cm; weight 17 kg (males) and 17 - 7 kg (females). Body measurements were taken from a sample in which the Terai Sacred Langur (S. hector) and the Nepal Sacred Langur were considered synonymous. The Nepal Sacred Langur, along with the Chamba Sacred Langur (S. ajax) and the Terai Sacred Langur, are reported to be the largest species of Semnopithecus, with some individuals weighing 20 kg or more. Coat of the Nepal Sacred Languris long and thick. It is sepia-brownish on back and outer limbs. Underside is white, and insides of limbs, rump, and tail tip are whitish. Whiskers around face are long and creamy. Nepal Sacred Langurs have bristle-like and lengthy eyebrows and a cream-hued crown, parted down the middle. Face is black.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFABFFAFFAEA6809F9F4F9F5.taxon	biology_ecology	Habitat. Subtropical to temperate broadleaved forest; semi-evergreen sal (Shorea robusta, Dipterocarpaceae) forest; alpine cedar forest; pine, riparian, and montane riparian forests; rocky outcrops; and scrub jungles in foothills of the Himalayas above elevations of 2000 m. Nepal Sacred Langurs have been observed at elevations as high as 3500 - 4000 m. Weather is highly seasonal, with harsh winters and rainy summers in July-October. Annual rainfall can be as high as 2500 mm.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFABFFAFFAEA6809F9F4F9F5.taxon	food_feeding	Food and Feeding. The diet of the Nepal Sacred Langur is made up of 45 - 60 % leaf parts (laminas, petioles, and buds) and 20 % ripe, unripe, and herbaceous fruits. It also contains seeds, roots, vegetative storage organs, flowers, bark, cambium, twigs, mosses, lichens, coniferous cones (Abies Pinaceae and Taxus Taxaceae), epiphytic fern rhizomes, grass, young bamboo shoots, and invertebrates. Nepal Sacred Langurs prefer broadleaved, deciduous leaves (both young and mature) over evergreen mature leaves. Seasonal variation in their feeding behavior is pronounced. They feed on leaf buds and ripe fruits of Cotoneasterfrigidus (Rosaceae), fruits of Berberis aristata (Berberidaceae), and leaf buds of Sorbus cuspidata (Rosaceae), herbs, and pinecones in winter. In spring and summer, their diets are more diverse. They feed on acorns, pinecones, young deciduous leaves, and young leaf clusters of Zanthoxylem nepalense (Rutaceae), and young leaves and bark ofJasminum humile (Oleaceae). During the monsoon, they feed mostly on young deciduous leaves and fruits. In autumn, they frequently consume unripe fruit (particularly legumes), seeds, and fleshy ripe fruits. They feed on soft underground fleshy plant tissues, herb fruits, and herb leaves. They feed extensively on cultivated potatoes Solanum tuberosum (Solanaceae). At higher elevations, they feed on pinecones, bark, and twigs.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFABFFAFFAEA6809F9F4F9F5.taxon	breeding	Breeding. Nepal Sacred Langurs have a birth season in late winter to early spring, allowing infants to grow sufficiently to survive their first winter in their montane habitats. Their independence is important for their mothers because they need to move long distances down slopes in the winter when it is snowing. Although Asian colobines, in general, wean their infants by the time they are one year old, Nepal Sacred Langurs have been found to have long interbirth intervals of c. 25 months and generally wean their young only after becoming pregnant again; lactation may last ¢. 26 months. Interbirth intervals are as short as 13 months if the previous infant dies. Mating increases in late spring (May). The menstrual cycle is not evident. Proceptive behavior by females is shown by presenting their anogenital region, with their tail lowered and simultaneously head-shaking. Copulation is always preceded by this behavior, but it does not always lead to mating. They mate throughout the year, and even pregnant females mate regularly, but sexual activity is highest in July-November, and the majority of conceptions (84 %) occur in July-September. Females maintain a dominance hierarchy in the group, and the dominant male interrupts mating attempts by subordinate males.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFABFFAFFAEA6809F9F4F9F5.taxon	activity	Activity patterns. Nepal Sacred Langurs are diurnal and mainly terrestrial. They prefer the upper forest canopy but also go to the ground to travel over rocks and through open areas close to forests. When fleeing, they prefer to move through trees rather than along the ground, as is typical of species in more open habitats. They sometimes sleep on semi-barren cliffs at elevations of ¢. 300 m or higher, taking advantage of their security and the radiant heat that rocks hold and the earlier sunrise that they receive.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFABFFAFFAEA6809F9F4F9F5.taxon	biology_ecology	Movements, Home range and Social organization. Nepal Sacred Langurs live in multimale-multifemale groups of 4 - 47 individuals, with 1 - 4 adult males and a male-to-female sex ratio of 1: 1 - 5 — 1 - 7. The meansize of five groups studied in 1972 - 1974 was 12 - 3 individuals. The same groups, and a sixth included in 1978, averaged 10 - 5 (resulting from a group splitting rather than a drop in population size). They also form all-male groups of ten or more, and males are sometimessolitary. Larger groups may split into dispersed subgroups, foraging separately while maintaining contact through intermittent calling by adult males. Annual home ranges of these temperate forest langurs are very large, 1275 - 12,700 ha, because they seasonally move up and down mountains. In mixed groups, there is a dominance hierarchy, and the dominant male frequently gives a deep contact call (" wao ” or “ ua ”), which even in the thick vegetation makes him the constant center of attention. Subordinate and subadult males usually travel on the periphery of the group. The Nepal Sacred Langur does not “ whoop-call ” in morning choruses as is typical of the species living on the plains further south (e. g. the Bengal Scared Langur, S. entellus, and the Tufted Gray Langur, S. priam). This may be because of the montane topography that renders such calling ineffective as a spacing mechanism. Female Bengal Sacred Langurs in conflict slap each other while sitting erect, face-to-face, but this has not been observed between female Nepal Sacred Langurs. Females have linear and stable dominance hierarchies, expressed by displacements. More dominant females maintain a better physical condition than lower ranking females. Infanticide following male takeovers has not been reported for Nepal Sacred Langurs, despite prolonged observations (32 months) in Junbesi-Ringmo between Kathmandu and the Nepal-Sikkim border.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFABFFAFFAEA6809F9F4F9F5.taxon	conservation	Status and Conservation. CITES Appendix I. Classified as Least Concern on The [IUCN Red List (2008). The Nepal Sacred Langur is protected under Schedule I, Part I, Indian Wildlife (Protection) Act, 1972, amended up to 2002, and the National Parks and Wildlife Conservation Act of 1973 in Nepal (listed as a common animal in Nepal). It is known to occur in at least ten protected areas: Chail Wildlife Santuary, Changthang Wildlife Sanctuary, Dachigam National Park, Hemis National Park, Kanchanjunga National Park, and Karakorum Wildlife Sanctuary in India; Mount Everest Natural Reserve, Langtang National Park, and Makalu Barun National Park in Nepal; and Manshi Reserve Forest in Pakistan. Major threats to Nepal Sacred Langurs include habitat loss; habitat degradation due to loss offruiting trees; timber, firewood, and charcoal extraction; forestfires; and encroachment by people. They are hunted and sold as a dry-meat delicacy in Nepal and used in native traditional medicine in China.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFAAFFA8FA3E6341FDD8F5F9.taxon	materials_examined	India, Travancore.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFAAFFA8FA3E6341FDD8F5F9.taxon	discussion	In his 1939 review of the Asian langurs, W. C. O. Hill listed six subspecies: hypoleucos, aeneas, lus, dussumaieri, achates, and elissa. C. P. Groves in 2001 considered aeneas to a be junior synonym, and dussumieri to be a distinct species, with ulus, achates, and elissa as synonoms. Groves’s research since that time has led him to believe that these forms (including dussumieri) require further study and that together they might result in at least one distinct subspecies of S. hypoleucos. M. L. Roonwal separated gray langurs of South Asia into a northern group and a southern group based on tail carriage. S. hypoleucos is of the southern group (Type IIA), with the tail held up and back, and the distal half to one-third dangling. Independent studies, based on mtDNA and nuclear DNA of the southern group of Hanuman Langurs, indicated that three subspecies are valid taxa. Three subspecies recognized.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFAAFFA8FA3E6341FDD8F5F9.taxon	distribution	Subspecies and Distribution. S. h. hypoleucosBlyth, 1841 — SWIndia (SWesternGhats), fromaroundBrahmagiriHillsinKarnatakaStateStotheNofSilentValleyinKeralaState. S. h. achatesPocock, 1928 — WIndia (WDeccanPlateauandEslopesofWesternGhats) uptotheRoonwallinethatfollowstheTaptiandGodavaririversintheN; limitedbythedistributionofS. entellusintheNEandS. priampriamintheE. S. h. wulus Pocock, 1928 — SW India (Western Ghats), from Jog Falls in Karnataka State, at 440 m, and S along the hilly wet zones to the Brahmagiri Hills.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFAAFFA8FA3E6341FDD8F5F9.taxon	description	Descriptive notes. Head-body 52 - 68: 6 cm, tail 73 - 109.2 cm; weight 13.6 - 15.9 kg (males) and 10.7 - 12.2 kg (females) for the “ Northern Malabar Langur ” (S. A. achates); head-body 50.8 - 55.5 cm, tail 72: 5 - 91.4 cm; weight 9 - 5 kg (males) and 8 - 4 kg (females) for the “ Black-legged Langur ” (S. A. iulus). Dorsum of the Malabar Sacred Langur is a deep mauve-brown, and undersides are pale orange. Limbs are black below elbows and knees. Reported exceptions include a population with pale upper arms. Yellowish-brown head has crown hairs that slope up at the front and smooth down at back end of head. Face and ears are black, and eyebrows are long and bristle-like. Tail is black, with a few white hairs at the tip.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFAAFFA8FA3E6341FDD8F5F9.taxon	biology_ecology	Habitat. Moist deciduous, riparian forests, sacred groves, and gardens at elevations of 100 - 1200 m. The Black-legged Langur and the “ Travancore Langur ” (S. A. hypoleucos) are specialized denizens of wet-evergreen and semi-evergreen montane forests. The Northern Malabar Langur occurs in drier areas and in plains from sea level to 700 m.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFAAFFA8FA3E6341FDD8F5F9.taxon	food_feeding	Food and Feeding. Malabar Sacred Langurs are folivorous, but when leaves are scarce, they eat flowers, buds, bark, and fruit. At Dharwar in the state of Karnataka, they spend 58 % of their time feeding on leaves, 29 % seeds and fruits, 7 % flowers, 2 % buds and bark, and 4 % other items such as insects. They eat leaf galls on Terminalia tomentosa (Combretaceae) and small black caterpillars. Water sources dry up in the dry season, but they can survive without drinking water for 4 - 5 months at a stretch.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFAAFFA8FA3E6341FDD8F5F9.taxon	breeding	Breeding. Newborn Malabar Sacred Langurs are seen in most months but with a somewhat higher frequency in December — April. In each group, births occur mainly within 2 - 6 months, but the months vary from group to group and from year to year. Infants are black and passed among females (allomothering). Males do not care for young, but the dominant male intervenes in defense of an infant when it is threatened (as it also does for any group member). Infants are weaned after ¢. 20 months, and even 2 year-old juveniles are sometimes seen suckling. Juveniles become completely independent of their mothers by three years of age. Infants are able to approach the dominant male, who tolerates and even plays with them.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFAAFFA8FA3E6341FDD8F5F9.taxon	activity	Activity patterns. Malabar Sacred Langurs are diurnal and mainly terrestrial. They spend ¢. 20 - 40 % of the day on the ground. When on the ground, they move about ten times faster and about three times farther than when in trees.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFAAFFA8FA3E6341FDD8F5F9.taxon	biology_ecology	Movements, Home range and Social organization. At Dharwar, the Northern Malabar Langur forms bisexual and all-male groups. Most bisexual groups have only one adult male (46 of 63 groups) and 6 - 9 adult females. Groups otherwise have 2 - 7 adult males, but many of them in this situation are believed to be undergoing a transition of male residence. Six bisexual groups studied at Dharwar were unimale — multifemale and contained 15 - 27 individuals. All-male groups contained 2 - 59 individuals. Bisexual groups have home ranges of 10 - 33 ha (average 18 - 6 ha). Home ranges of all-male groups overlap those of several bisexual groups. Adult males disperse and form allmale groups. Contacts among bisexual groups and all-male groups are generally aggressive and have been described as attacks. When the dominant male of a bisexual group sees an all-male group, he approaches and threatens them (grinding his teeth, biting the air, and display jumping, accompanied by “ whoops ” and belching). Often, the all-male group runs away, but sometimes males counterattack, resulting in flight or injury of the bisexual group’s male, and then the all-male group invades his group. Sometimes invading males leave the area after a few hours, and the group’s male, if not too injured, returns. Sometimes males that carry out a successful attack on a bisexual group drive out the resident adult, subadult, and sometimes juvenile males, or abduct the group’s adult and immature females, while killing the infants. The majority of attacks occur during the first half of the mating season. Population densities at Dharwar were very high at 85 - 134 ind / km * when Y. Sugiyama and K. Yoshiba carried out their studies there. All-male groups were common, and the frequency of social changes in bisexual groups was high. The two longest recorded male tenures at Dharwar were three and six years. The high densities possibly resulted from lack of predators and loss of habitat, concentrating individuals into a small area.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFAAFFA8FA3E6341FDD8F5F9.taxon	conservation	Status and Conservation. CITES Appendix I. Classified as Vulnerable on The IUCN Red List, including the nominate subspecies, but the subspecies achates (under S. dussumier: achates) and ulus (under S. dussumieri iulus) are classified as Least Concern. The Malabar Sacred Langur is protected under Schedule II, Part II, of the Indian (Wildlife) Protection Act, 1972, amended up to 2002. The Travancore Langur occurs in Wyanad and Brahmagiri wildlife sanctuaries and Nagarhole National Park. The Black-legged Langur occurs in Kudremukh, Mollem, and Nagarhole national parks and Pushpagirl, Bhadra, Sharavati Valley, Anshi, Bondla, Mookambika, and Radhanagari wildlife sanctuaries. Hunting is a serious threat, particularly for the two western hill zone subspecies, the Travancore and Black-legged langurs. The Malabar Sacred Langur has been entirely extirpated from the eastern slopes of Coorg because of hunting. Other threats include fragmentation, habitat loss, local trade in live animals and meat, trade for use in traditional medicine, developmental activities, and change in land use for agriculture and mining. The Malabar Sacred Langur is the most poorly studied of the southern Indian langurs.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFADFFA9FFEE6F59FF00FA21.taxon	materials_examined	India, Coromandel Coast.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFADFFA9FFEE6F59FF00FA21.taxon	discussion	M. L. Roonwal separated gray langurs of South Asia into a northern group and a southern group based on tail carriage. S. priam is of the southern group (Type IIB), with the tail held up in a letter “ S, ” and the distal one-third pointing backward, not dangling. In his 1939 review of the Asian langurs, W. C. O. Hill considered the forms anchises and thersites to be subspecies of S. priam, and recent molecular studies based on mtDNA and nuclear DNA have confirmed this. Three subspecies recognized.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFADFFA9FFEE6F59FF00FA21.taxon	distribution	Subspecies and Distribution. S. p. priam Blyth, 1844 — S & SE India (Andhra Pradesh, Karnataka, Kerala, and Tamil Nadu states), a highly fragmented distribution ranging from the Krishna River in Andhra Pradesh S to Tirunelveli in Tamil Nadu. S. p. anchises Blyth 1844 — SC India (S Deccan Plateau), found in the districts of Kurnool, Andhra Pradesh, and in Pavagada in the district of Tumkur, Karnataka. S. p. thersites Blyth 1847 — Dry Zone of Sri Lanka, ranging from Jaffna in the N to the S coast.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFADFFA9FFEE6F59FF00FA21.taxon	description	Descriptive notes. Head-body 58.4 - 64 cm, tail 66 - 101.6 cm; weight 11.4 - 13.5 kg (males). The Tufted Gray Langur is the smallest of the gray langurs. In the “ Central Indian Gray Langur ” (S. p. anchises), coat is brownish gray, variable with some individuals having darker coats in wetter areas and duller hues in drier areas. Its belly and chest have an orange tinge, well pronounced and prominent in wetter coastal habitats and duller, paler in drier areas. It has no crest. Its palms are dark up to knuckles. The “ Madras Gray Langur ” (S. p. priam) has a conspicuous coconut tuft-like crest on its head. Its fur is grayish-brown on crown, dorsum, arms, upper thighs, and tail. It is creamy white on head, including cheek whiskers, and white on rump. Its limbs are paler than its body and either gray, blond, or near-white. Skin on its face and ears is black. It has long and bristle-like eyebrows. The “ Sri Lankan Gray Langur ” (S. p. thersites) has creamy white cheek and chin whiskers, a distinctive pointed, brown-coconut tuft-like crest on its crown, and a prominent fringe. It is brownish-gray (darker than the Madras Gray Langur) on its dorsum, and paler on its ventrum, hands, and limbs. Its tail is dark, with a white tip.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFADFFA9FFEE6F59FF00FA21.taxon	biology_ecology	Habitat. Mostly in forested areas, occasionally in home gardens, at elevations of 180 — 1200 m. The “ Central Indian Gray Langur ” (S. p. anchises) is found in tropical dry deciduous forests, occupying the middle to upper canopy at elevations up to 800 m, as well as near human settlements and forest fringes. The Madras Gray Langur is found at elevations of 180 - 1200 m in highly degraded, non-contiguous tropical dry evergreen forest and scrub forest. The Sri Lankan Gray Langur, endemic to Sri Lanka, is found in dry, deciduous forests in lowlands, and it overlaps with the Purple-faced Langur (S. vetulus) in semi-evergreen forests. It does not occur in wetter evergreen forests and hill zones at elevations above 500 m in southern Sri Lanka, which is the exclusive domain of the Purple-faced Langur. It can also be found in urban outskirts and temples. The Sri Lankan Dry Zone has a marked dry season in June — October and annual rainfall is ¢. 1700 mm. Sri Lankan Gray Langurs inhabit alluvial and high stature forest (15 m high or more) and medium and low stature forest (5 - 15 m), low scrub, and slash-and-burn cultivation.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFADFFA9FFEE6F59FF00FA21.taxon	food_feeding	Food and Feeding. Most of what is known about the Tufted Gray Langur comes from studies of the Sri Lankan Gray Langur in the Sri Lankan Dry Zone at Polonnaruwa. They spent 48 % of their feeding time eating leaves and shoots (less than 50 % of this on mature leaves), 45 % fruits and seeds (generally mature and fleshy fruits), and 7 % flowers. They also ate termite clay. They were more frugivorous, living in larger groups, and using larger home ranges than the partially sympatric Purple-faced Langur. In some months, fruits made up 80 - 90 % of the diet. In any one month, mature leaves never made up more than 50 % of the diet. In this study, 90 % of the diet was supplied by 23 tree species; 30 % by three species — Walsura piscidia (Meliacaeae), Drypetes sepiaria (Euphorbiaceae), and Schleichera oleosa (Sapindacae) — and 50 % by five species — adding Adina cordifolia (Rubiaceae) and Ficus benghalensis (Moraceae). Another three Ficus species were also included in the diet. During the monsoon in February-March, they ate mainly young leaves and shoots, especially of Walsura and Drypetes, and smaller amounts of mature leaves. In the dry season in May — October, they were more frugivorous; fruits were more abundant then, especially Drypetes and Ficus, and fruits dominated the diet in April-September. Flowers were eaten mainly during the dry season, and leaves were again the principal items in the second monsoon in October-November and rains in December — January. When near settlements and villages, Sri Lankan Gray Langurs also feed on fruits, flowers, and cultivated crops. In some areas people feed them, and this can be a significant portion of their diet.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFADFFA9FFEE6F59FF00FA21.taxon	breeding	Breeding. Although little is known about breeding patterns of the Tufted Gray Langur, it is likely similar to its congener, the Malabar Sacred Langur (S. hypoleucos). Births occur throughout the year. Females have a 24 day menstrual cycle and are receptive through the year. The gestation period is 168 - 200 days, and the interbirth interval is 16 - 7 months. Infants are weaned after 13 - 20 months.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFADFFA9FFEE6F59FF00FA21.taxon	activity	Activity patterns. The Tufted Gray Langur is diurnal and mainly terrestrial. Foraging activity peaks in early morning at 06: 00 - 09: 30 h and again before nightfall at 15: 30 - 18: 00 h. Intensive feeding takes up to c. 3 hours / day.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFADFFA9FFEE6F59FF00FA21.taxon	biology_ecology	Movements, Home range and Social organization. There is no definitive information for the mainland populations of Tufted Gray Langurs. The Sri Lankan Gray Langur lives in unimale — multifemale and multimale-multifemale groups of 20 - 60 individuals, with 3 - 6 adult males. They live in home ranges of 30 - 130 ha that overlap extensively. Core areas of exclusive use are 7 - 20 ha. Group sizes and home ranges tend to be larger in open habitats than in forests. At Yala National Park in south-eastern Sri Lanka, groups have 40 - 60 individuals, with 8 - 10 adult males. It would seem that even in these dry areas, Tufted Gray Langurs are not very dependent on free water. A group studied at Polonnurawa lived near a reservoir but rarely drank from it. Considered to be a spacing mechanism among groups, males give loud “ whooping calls ” in choruses in early to mid-morning (after sympatric Purple-faced Langurs, which generally call before sunrise). Sometimes Tufted Gray Langurs go to the tops of emergent tamarind trees (Tamanindus indica, Fabaceae) for these choruses. A distinctive feature of their social behavior is the “ display jump, ” performed at any time during the day, usually stimulated by hearing another group. Males whoop and jump between branches, often, it seems, breaking them intentionally, in a heavy stiff fashion. They are sometimes accompanied in these displays by females without young. The Tufted Gray Languris largely terrestrial (c. 80 % of time). They move quadrupedally or with a leaping gait through the forest. When not eating, females groom, tend their infants, or rest, whereas males tend to sit up in a tree, giving low calls (especially grunts), watching the group, and presumably being vigilant for other groups or predators. Group encounters are quite frequent (about one every ten hours) and are generally antagonistic, involving chases, displacements, threats, mild wrestling, grabbing, and sometimes fighting. It is evident that groups force these encounters in that they travel far off their habitual routes to find their neighbors. Interactions of this sort can last up to two hours. Group takeovers are common, with new males aggressively ousting the existing alpha male. Infants and Juveniles are often killed in such troop takeovers.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFADFFA9FFEE6F59FF00FA21.taxon	conservation	Status and Conservation. CITES Appendix I. Classified as Near Threatened on The IUCN Red List, including the nominate subspecies priam. The subspecies thersites is classified as Endangered and anchises (under S. dussumieri anchises) as Least Concern on The IUCN Red List. It is protected under Schedule II, Part I, of the Indian Wildlife (Protection) Act, 1972. Primary threats to Tufted Gray Langurs are habitat loss, wildfires, hunting for food, and conflict with people (crop raiding). The Central Indian Gray Langur occurs in the Nagarjunasagar-Srisailam Tiger Reserve. The Madras Gray Langur is declining in Andhra Pradesh because of hunting and habitat loss. Hunting is rampant in the Eastern Ghats and along the eastern coast. Tufted Gray Langurs occur in nine protected areas in India: Bandipur and Sri Venkateshwara national parks; Cauvery, Biligiri Rangaswamy Temple, Mudumalai, Indira Gandhi, Chinnar, and Nellapattu wildlife sanctuaries; and Kalakkad-Mundanthurai Tiger Reserve. The Sri Lankan Gray Langur is protected under the Fauna and Flora Protection Ordinance Act No. 2, 1937, and its subsequent amendments, including Act No. 49, 1993, in Sri Lanka. It occurs in Ritigala Strict Nature Reserve; Victoria-Randenigala-Rantambe; Knuckles Forest; Ampara, Buddaragala and Kanthale Naval wildlife sanctuaries; and Wilpattu, Angamedilla, Flood Plains, Giritale, Moragaswawe, Somawathie, Wasagamuwa, Udawalawe, Bundala, Lunugamvehara, Madura Oya, and Ruhuna national parks.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFACFFAAFF286336FBE9F907.taxon	vernacular_names	Purple-faced Langur	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFACFFAAFF286336FBE9F907.taxon	vernacular_names	French: Langur blanchatre / German: Weibartlangur / Spanish: Langur purpura Other common names: Purple-faced Leaf Monkey, Purple-faced Monkey; Bear Monkey / Highland Purple-faced Langur / Montane Purple-faced Langur (monticola), Dry Zone Purple-faced Langur / Northern Purple-faced Langur (philbricki), North Lowland Wet Zone Purple-faced Langur / Western Purple-faced Langur (nestor), Southern Lowland Wet Zone Purple-faced Langur / Southern Purple-faced Langur (vetulus)	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFACFFAAFF286336FBE9F907.taxon	materials_examined	Ceylon (= Sri Lanka).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFACFFAAFF286336FBE9F907.taxon	discussion	Molecular data support the classification of the gray or Hanuman langurs, S. vetulus, and S. johnii, in the same genus, Semnopithecus. Despite its restricted geographic distribution, S. wvetulus has four well-recognized subspecies. The subspecies were recognized by W. C. O. Hill in his 1939 re-view of the Asian langurs. They are quite distinct and further study may indicate that they should each be considered species. An additional subspecies, harti, described by Deraniyagala in 1955, is also recognized by some experts. It is said to be like the form philbricki but darker, more reddish-brown and with a more prominent light rump patch. Four subspecies recognized.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFACFFAAFF286336FBE9F907.taxon	distribution	Subspecies and Distribution. S. p. vetulusErxleben, 1777 — SWSriLanka (WetZone), fromKaluRiverStoRama, toanelevationof 1000 m; thedistributionislessthan 5000 km?. S. p. monticolaKelaart, 1850 — MtsofCSriLankaatelevationsof 1000 - 2200 m. S. p. nestorBennett, 1833 — WSriLanka (WetZone), NoftheKaluRiveratelevationsupto 1000 m. S. p. philbricki Phillips, 1927 — N & E Sri Lanka (Dry Zone) up to elevations of 1500 m in E Matale and Madulkele Hills.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFACFFAAFF286336FBE9F907.taxon	description	Descriptive notes. Head-body 50 - 65 cm (males) and 45 - 60 cm (females), tail 67 — 85 cm (males) and 62 - 82 cm (females); weight 3.4 - 9.4 kg (males) and 3.8 - 9.3 kg (females). The Purple-faced Languris generally brown or blackish-brown, with a whitish throat, interramal region (under the chin), and tail. Tail has a gray or white triangular patch at its root. Face is purplish-black (not actually purple) and lined with a ridge of black fur on forehead. In adults, there are long, pale, backswept, white (sometimes brown-tipped) cheek whiskers, contrasting with a brownish crown. Females have a white pubic patch. The “ Southern Purple-faced Langur ” (S. v. vetulus) is small and sexually dimorphic. Its fur is generally grayish-black, with whitish-brown on lower back, haunches, and distal part oftail. Its head is brown or grayish-brown, contrasting with its body. Its cheek whiskers are also grayish-brown and only cover lower half of ears. The “ Western Purple-faced Langur ” (S. v. nestor) is a somewhat small, pale form, with little or no sexual dimorphism. Its back and sides are predominantly grayish-brown, rump is silvery-gray, forearms and shanks are nearly black, and crown and nape are pale brown. Its tail is slightly tufted at its tip. The “ Northern Purple-faced Langur ” (S. v. philbricki) is a brown form, very like the Western Purple-faced Langur, exceptit has a less conspicuous rump patch. Lower parts of limbs are black, and tail tip is pale tawny or white and untufted. Its cheek whiskers are very long. The “ Highland Purple-faced Langur ” (S. v. monticola) is a large, robust, montane subspecies. Its fur is thick and brown, with only slightly contrasting crown and rump patch. Its cheek whiskers are long and white, and they hide its ears. Its tail is notably short. A population of the Southern Purple-faced Langur was recently discovered in the districts of Galle and Matara in far south-western Sri Lanka that has extraordinary and beautiful variations in pelage color and patterns, with different extents and patterns of white; they are collectively called “ white-color morphs. ”	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFACFFAAFF286336FBE9F907.taxon	biology_ecology	Habitat. Wet forests, parkland, and dense primary montane forests to elevations of 2200 m. The Southern Purple-faced Langur is found in low and middle elevation rainforests, plantations, and home gardens with adequate canopy cover. The Western Purple-faced Langur is found in low-elevation tropical rainforest, with refugee populations in semi-urban and rural home gardens, rubber plantations, and any areas with groves of trees and adequate canopy cover. The Highland Purple-faced Langur occurs in montane tropical rainforest. The Northern Purple-faced Langur occurs in dry evergreen and deciduous forests, restricted to moister areas with closed canopy and permanent water sources in dry-zone forests. In the southern part ofits distribution near Galle, the Southern Purple-faced Langur forages on rocky, treeless coastal slopes.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFACFFAAFF286336FBE9F907.taxon	food_feeding	Food and Feeding. The Purple-faced Langur eats leaves (young and mature), shoots, fruits, seeds, and flowers, and occasionally bark and termite nest soil. Its diet is not very diverse, and they tend to exploit a small number of common speciesin relatively small home ranges. In terms of the time spent feeding, the diet of the Northern Purplefaced Langur at Polonnaruwa contained 40 % mature leaves, 28 % seeds and fruits, 20 % young leaves and shoots, and 12 % flowers. Most ofits food (more than 90 %) was taken from only about twelve plant species. For one group, 40 % of the feeding time was dedicated to leaves, flowers, and fruits of Adina cordifolia (Rubicaeae); two other species, Schileichera oleosa (Sapindaceae) and Drypetes sepiaria (Euphorbiaceae), made up to 30 % of the diet. A brief study of the diet of the Highland Purple-faced Langur (25 hours of contact) indicated that it was more dependent on young leaves and shoots, with a diet of 75 % young leaves, 16 % coriaceous (thick, leathery) leaves, and 10 % flowers and fruits. Leaves, especially young leaves and shoots, account for more than 95 % of the diet of the Northern Purple-faced Langur during the February — - March monsoon. Fruits make up a higher portion of the diet in the dry season, including unripe fruits of Elaeodendron glaucum (Celastraceae), and flowers of Adina cordifolia (Rubiaceae) are also eaten during the dry months in July-September. The Purple-faced Langur is consistently more folivorous, eating more leaves, than the sympatric Sri Lankan Gray Langur (S. priam thersites) that eats more shoots and fruits. In human-modified environments, the Western Purple-faced Langur exploits cultivated fruits, particularly jackfruit (Artocarpus heterophyllus, Moraceae), mango (Mangifera indica, Anacardiacae), dehiscent seeds of rubber (Hevea brasiliensis, Euphorbiaceae), and bananas. They occasionally raid potato and cauliflower crops. Two groups of Western Purple-faced Langurs, living around villages in home ranges that had small groves of trees, plantations, and gardens, had diets that contained a little more than 50 % fruit, principallyjackfruit, 16 - 5 - 22 % young leaves, and 8 - 13 % mature leaves and leaf stems.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFACFFAAFF286336FBE9F907.taxon	breeding	Breeding. Breeding of Purple-faced Langurs can take place at any time of the year, although mating of the Southern Purple-faced Langur occurs mainly during the rainy season in October — January when food is abundant, temperatures are falling, and day length is shortening. Births peak in the dry season. Climatic variation is less marked on the Horton Plains, so the Highland Purple-faced Langur does not have a birth peak. A single young is born after a gestation of 195 - 210 days. The Highland Purple-faced Langur has an interbirth interval of 16 - 17 months, whereas the Southern Purple-faced Langur in the more seasonal climate at Polonnaruwa has an interbirth of ¢. 24 months. Infants under four months of age are pale gray, with a brownish tinge on their chests, arms, legs, and crowns; naked parts of their skin are black and their cheek hairs white. Allomothering by females is common; males tend to ignore infants. Weaning begins when infants are 7 - 8 months old. Infanticide by adult males has been reported.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFACFFAAFF286336FBE9F907.taxon	activity	Activity patterns. Purple-faced Langurs are diurnal and, unlike gray langurs, largely arboreal.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFACFFAAFF286336FBE9F907.taxon	biology_ecology	Movements, Home range and Social organization. Purple-faced Langurs generally live in small unimale-multifemale groups that average 8 - 4 individuals, with 3 - 5 females and their offspring. Multimale-multifemale groups have been observed in the Western Purple-faced Langur. Groups as large as 20 individuals have been recorded, butrecent (2007) range-wide surveys of the Western Purple-faced Langur found that groups were small in many areas, averaging 4 - 4 (range 1 - 10, n = 9) — believed to result from deforestation and hunting. Unimale-multifemale groups of Highland Purple-faced Langur average 8 - 9 (range 3 - 14, n = 27), and those of the northern and eastern subspecies, the Northern Purple-faced Langur, average 8 - 4 individuals (range 3 - 14, n = 33). They occupy small home ranges, and males are very territorial, giving spectacular displays of dropping leaps accompanied by loud harsh calls and “ whooping ” to advertise their residency when they meet other males. Home range sizes are 0 - 9 - 8 - 6 ha (average 3 - 1 ha) for bisexual groups of Southern Purple-faced Langurs and 1 - 1 - 10 - 9 ha (average 5 - 8 ha) for Highland Purple-faced Langurs. Two groups of Western Purplefaced Langurs in semi-urban landscapes occupied areas of 3 - 9 ha and 9 - 6 ha. Males disperse and live in predominantly all-male groups. Male group takeovers are violent, and a successful male expels resident males and subadults and juveniles and kills the infants. Subadults and juveniles join the predominantly male groups, which are nonreproductive and notat all cohesive; they generally occupy marginal habitats, degraded with low and disrupted canopies. In one case, a female with an older infant left the group when a new male entered and stayed with the deposed male until her infant was fully grown. Male tenure of a heterosexual group lasts c. 3 years. Females will mate with subadult males, which is largely tolerated by the alpha male — presumably such matings are non-conceptive.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFACFFAAFF286336FBE9F907.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Endangered on The [UCN Red List (as Trachypithecus vetulus), including the subspecies vetulus, philbricki, and monticola, but the subspecies nestor is classified as Critically Endangered. The Purple-faced Langur is threatened by hunting and is particularly sensitive to deforestation. The Western Purple-faced Langur has been severely affected by the almost total loss of forest in the Sri Lankan Wet Zone. All subspecies have small distributions and have suffered major declines in numbers over the last 20 years through the fragmentation and degradation of their forests. In the absence of hunting, however, their ability to adapt to human-modified landscapes is astonishing. They make use of electricity pylons, powerlines (electrocution is frequent), roof tops, and fences in mosaics of tea plantations, rice paddy, rubber plantations, cinnamon and cardamom plots, and home gardens to move through their home ranges and to find food and sleeping sites. Sometimes Purple-faced Langurs are forced to use the ground and run the risk of being killed by dogs or caughtto be reared as pets. Antagonism toward Purple-faced Langurs is high when they raid home gardens and destroy rooftiles. Feeding on garbage is also a threat because food waste is undoubtedly unhealthy for them and a means of disease transmission. Surveys have shown that more than 50 % of the population of Western Purple-faced Langurs now comprisesisolated pockets of one or two groups in matrices of urbanized landscapes, village settlements, fields, and plantations. Their crop raiding means that conflict with humansis prevalent, even though they are generally not hunted. Thissituation cries out for proactive management and conservation measures such as improving the management of existing protected areas, creating new protected areas, and mitigating the discomfort of humans and langurs when they are in such close and constant contact. Purple-faced Langurs occur in a number of protected areas including Angamedilla, Flood Plains, Horton Plains, Kaudulla, Madura Oya, Minneriya-Giritale, Moragasawe, Somawathie, Udawalawe, Wasgamova and Wilpattu national parks; Anuradhapura, Kitulgala, Kurulukale, Mihintale, Peak Wilderness, Polonnaruwa, and Rantembe sanctuaries; Hakgala and Ritigala nature reserves; and Dombaghakanda and Knuckles forest reserves. Ingiriya Forest Reserve is an important refuge for the Western Purple-faced Langur. The two largest forests remaining in the distribution of the Purple-faced Langur surround two reservoirs, Kalatuawa and Labugama, which supply water to Colombo, the country’s capital. These forests encompass 2100 ha, part of which have already been turned into pine plantations. Preservation of these forests should be a very high priority in any conservation strategy on behalf of the Purple-faced Langur.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFAFFFABFFEE6254F97AFE3E.taxon	materials_examined	India, Tellicherry.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFAFFFABFFEE6254F97AFE3E.taxon	discussion	Molecular data support the classification of the gray or Hanuman langurs, S. vetulus, and S. johnii, in the same genus, Semnopithecus. There is some apparent geographic variation in the presence and extent of silvering on the rump. In the northern part of the distribution, it is quite conspicuous, even extending forward onto the back or backward along the tail. S. johnii is believed to hybridize with S. hypoleucos where sympatric. Such crosses tend to be brownish. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFAFFFABFFEE6254F97AFE3E.taxon	distribution	Distribution. SW India (Karnataka, Kerala, and Tamil Nadu states); it occurs sporadically in montane forests of the Western Ghats from Srimangala (12 ° 01 ’ N, 75 ° 58 ’ E) in Karnataka S to the Aramboli Pass (8 ° 16 ’ N) in Kerala.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFAFFFABFFEE6254F97AFE3E.taxon	description	Descriptive notes. Head-body ¢. 78 cm (males) and c. 58 - 4 cm (females), tail 68.5 - 96 cm; weight 11.8 - 13.6 kg (males) and 9.8 - 11.3 kg (females). Coat of the Nilgiri Langur consists of glossy black hairs with brown bases. Old individuals have white hair on back and rump. Face is black to purplish-red, and crown, sides of head, and cheeks are a light brown to vivid brownish-orange, becoming yellowish-brown on nape and sometimes on brow; contrast between these bright and dark areas is quite startling and is further accentuated by a narrow black brow fringe. Crown hairs are posteriorly directed. Tail is black, although sometimes partly silvery-gray at the base, the color extending to the rump. Sexes are similar, but females are alone in having small patches of white fur on insides of their thighs, which are present even in very young individuals. Males have larger canines than females. Differences in coat color have been recorded at Agastya Mali, the most southerly hill range in the Western Ghats. Specifically, they were in the Kalakkadu and Mundanthurai wildlife sanctuaries in Tamil Nadu, on the east side of the distribution. Groups there included individuals (more than one year old) with pale chocolate-brown coats and golden or silvery-golden hairs on their head. They were in groups that otherwise contained Nilgiri Langurs with typical black coats. They were seen in an area where langur groups were either completely isolated or abutted the home range of just one other group, indicating it may be a founder effect (if they were colonizing) or caused by inbreeding (if they had been isolated by deforestation).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFAFFFABFFEE6254F97AFE3E.taxon	biology_ecology	Habitat. Evergreen, semi-evergreen, montane, and riparian forests; “ sholas ” (narrow tracts of evergreen forest, surrounded by grassland, usually with a stream running through them); and moist deciduous forests of the Western Ghats at elevations of 300 - 2400 m. Nilgiri Langurs are also seen in teak plantations close to natural forests.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFAFFFABFFEE6254F97AFE3E.taxon	food_feeding	Food and Feeding. Nilgiri Langurs feed on leaves (both young and mature), fruits, flowers, seeds, bark, insects, and termite soil. More than 100 plant species have been identified in their diet, but ¢. 50 % of their feeding time tends to be dedicated to a few species (three in one study, ten in another). In a study of two groups over 24 months in lowland riparian forest on the Mundanthurai Plateau, their feeding time was apportioned as follows: 44 % young leaves, 19 % seeds, 10 % unripe fruits, 8 % flowers, 5 % ripe fruit, 4 % mature leaves, and 10 % other food items such as petioles, bark, pith, insects, gum, and wood. They fed on fresh ripe fruits during monsoon and post-monsoon seasons and gradually shifted to seeds with the onset of the dry season. Although they ate mature leaves, seasonally available young foliage, flowers, and fruits were preferred. In another study of a group of Nilgiri Langurs in tall, closed canopy, evergreen forest at Kakachi in the Agastya Malia range, the plant part of the diet included 25 - 5 % young leaves, 25 - 1 % fruits and seeds, 20 - 3 % mature leaves (mostly from trees), 9 - 3 % flowers and flower buds, 6 - 5 % petioles of mature leaves, 5: 7 % leaf buds and initial flush of foliage, 4 - 2 % leaves of unknown age, and 3.4 % stems, bark, and undetermined items. In this study, young foliage was eaten from 56 species, and mature foliage was taken from only 39. Of 115 plant species in the diet, 58 were trees (including a tree fern), 32 were climbing plants, 13 were non-woody herbs, six were woody shrubs and bamboos, and six were parasites or epiphytes. Strong selectivity in their diet was demonstrated in that 45 % of the diet came from only three tree species and the two most common trees in the forest contributed only 5 %. Nilgiri Langurs also occasionally eat insects, including termites and stick insects. They raid potato, cauliflower, cabbage, and cardamom crops.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFAFFFABFFEE6254F97AFE3E.taxon	breeding	Breeding. Female Nilgiri Langurs do not show any known physical or behavioralsigns of sexual receptivity, although they have an everted pink clitoris that changes color, darkening at times and possibly associated with the periovulatory period of the menstrual cycle. A single young is born every 1 - 5 - 2 years, usually in May-November. Infants are born with pale skin and reddish-brown body hair, which gradually darkens and is close to adult coloration by 4 - 5 months of age. Mothers carry their young on their belly. Allomothering has been reported. Birth rates of groups in low-elevation riverine habitats are lower than those living at higher elevations (0 - 2 vs. 0 - 7 births per female / year).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFAFFFABFFEE6254F97AFE3E.taxon	activity	Activity patterns. Nilgiri Langurs are diurnal, arboreal, and infrequently terrestrial. They are known to move to moist deciduous forests in morning and return to evergreen forests at dusk. They move in single file, using the same route from branch to branch as their leader, usually in the middle forest canopy. Peak feeding time is in the morning and evening, and midday is rest time. They are generally found in the understory at heights of 5 - 9 m, but they sleep, feed, and sun themselves in the upper canopy at 12 - 24 m. They rarely go to the ground where they sometimes feed, fight, and play.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFAFFFABFFEE6254F97AFE3E.taxon	biology_ecology	Movements, Home range and Social organization. The Nilgiri Langur typically lives in unimale-multifemale groups of ¢. 7 - 9 individuals, but group sizes can vary from three to 27 individuals. Female emigration from the natal group is common and is a strategy to minimize competition and infanticide and increase access to resources. Dispersing males generally remain solitary until they can enter a group and take over as the dominant male. In one site where Nilgiri Langurs have been studied (Ootacamund), they also form small, ephemeral predominantly male, non-reproductive groups. Rarely, they form all-female groups. Sometimes new groups are formed by splitting — unusual among langurs. Home ranges vary in size (5 - 260 ha), evidently influenced by group size and dispersion, and the abundance and seasonality of foods in the different forest types. Home ranges of groups living in sholas tend to be larger. There is little overlap, and they are actively defended by adult males. When facing off with each other, males typically sit on high branches, open their mouths and emit continuous low buzzes that sound like creaking doors, and snap their heads upward with their mouths open, appearing to bite the air. Chases are accompanied by “ whoops, ” grunts, hiccups, and “ hah-hah ” calls. Group spacing is maintained by whooping sessions and vigilance by males. The adult male in a unimale-multifemale group rarely interacts with females. Females sometimes associate with Malabar Sacred Langurs (S. hypoleucos) and Bonnet Macaques (Macaca radiata) until they form a heterosexual (one-male) group. When not hunted, Nilgiri Langurs reach high densities of 70 ind / km?.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFAFFFABFFEE6254F97AFE3E.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Vulnerable on The IUCN Red List (as Trachypithecus johnii). The Nilgiri Langur is included in Schedule I, Part I of the Indian Wildlife (Protection) Act, 1972, amended up to 2002. Despite its full legal protection, it is still hunted because its flesh is considered highly flavorful. Its skin is used to make ornamental drums. It is also much sought after by poachers for its blood and internal organs, which are widely believed to cure variousliver and lung ailments and to act as an aphrodisiac (flesh and glands are used to “ treat ” asthma, and the blood is drunk fresh by locals as a “ rejuvenator ”). Body parts are used to make medicinal potions called “ Karum Kurangu Rasayanam ” (Tamil for “ Langur tonic ”). These direct pressures on Nilgiri Langurs are exacerbated by the steady loss, degradation, and fragmentation of their already limited habitat, as a result of mining, dam construction, and encroachment of plantations. As of 2003, the total wild population was estimated at less than 10,000 mature individuals. Nilgiri Langurs occupy a mere 16 % of the total area of the Western Ghats. Areas surrounding sanctuaries and national parks suffer from anthropogenic pressures (extraction of timber and non-timber resources), and most tree species that are valued for their timber are also part ofthe preferred diet of Nilgiri Langurs. Domestic dogs prey on Nilgiri Langurs when they move between forest patches. Nilgiri Langurs occur in Eravikulam, Mukurthi, Periyar, and Silent Valley national parks and Aaralam, Brahmagiri, Chimmony, Chinnar, Grizzled Giant Squirrel, Idukki, Indira Gandhi, Kakkal, Kalakkad, Mudumalai, Mundanthurai, Neyyar, Parambikulam, Peechi, Peppara, Shendurney, Thattekadu, and Wyanad wildlife sanctuaries.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB0FFB5FF2564A7FC60F3C9.taxon	materials_examined	India, Assam, Goalpara district, Jamduar Forest Rest House, east bank of Sankosh River.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB0FFB5FF2564A7FC60F3C9.taxon	discussion	T. geei forms, together with 1. pileatus and T. shortridgei, the pileatus species group. The phylogenetic position of this group remains unresolved, and they might have evolved through hybridization of ancestors (3: 4 - 7 - 1 million years ago) of today’s genera Trachypithecus and Semnopithecus. A subspecies, bhutanensis, was described in 2003, but the possibility remains thatit is from a population of recent hybrids, and it is not recognized here. 1. geei is marginally sympatric with T. pileatus in a small area of Bongaigaon district, Assam. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB0FFB5FF2564A7FC60F3C9.taxon	distribution	Distribution. NE India (Himalayan foothills of NW Assam State between the Manas and Sankosh rivers, and in the area immediately to the S, nearly to the Brahmaputra River) and Bhutan (as far N as Black Mt, ¢. 27 ° 30 ’ N, E of the Sankosh River; also extends E of the Mangde River along either side of the main stream).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB0FFB5FF2564A7FC60F3C9.taxon	description	Descriptive notes. Head-body 64 - 72 cm (males) and 48.8 - 61 cm (females), tail 78 94 cm (males) and 71 - 80.5 cm (females); weight 10 - 12 kg (males) and 9.5 - 10 kg (females). Coat of the Golden Langur is normally a uniform vivid flaxen-yellow to creamy-white and pelage is notably long and lax. There are long, backward-swept cheek whiskers that extend to ears and an ill-defined whorl of dark hairs across forehead. Face is naked and black. An external scrotum is absent. Coat coloris reported to take on a reddish tinge in summer and a whitish tinge in winter.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB0FFB5FF2564A7FC60F3C9.taxon	biology_ecology	Habitat. Evergreen, tall sal (Shorea robusta, Dipterocarpacae) moist deciduous, riparlan, savanna, temperate, and subalpine forests, up to elevations of 3000 m in the northern part of its distribution. The Golden Languris often found in plantations. Tall (up to 45 m) moist deciduous forests have leafless canopies during the dry season. At the end of the dry season, there are flushes of new leaves and flowers.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB0FFB5FF2564A7FC60F3C9.taxon	food_feeding	Food and Feeding. Golden Langurs eat mainly young leaves and buds, along with mature leaves, fruits (ripe and unripe), seeds, and flowers. Young leaves of Terminalia belerica and I. crenulata (Combretaceae), buds and flowers of Dillenia pentagyna (Dilleniaceae), Careya arborea (Lecythidaceae), and Bombax ceiba (Malvaceae), and leaves of Bauhinia vahli (Fabaceae) are important foods. Golden Langurs eat soil at clay licks. They raid cardamom, tapioca, and guava crops.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB0FFB5FF2564A7FC60F3C9.taxon	breeding	Breeding. Mating season of the Golden Languris inJanuary-February. A single offspring is born in July-August, after a 6 month gestation. Infants are orange-brown or gray.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB0FFB5FF2564A7FC60F3C9.taxon	activity	Activity patterns. Golden Langurs are diurnal and arboreal. They are active after dawn and feed mostly in early morning until ¢. 09: 30 h and again in late afternoon at c. 16: 00 - 17: 00 h. They rest in the middle of the day in the tops oftall trees and huddle in the shade under vine tangles.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB0FFB5FF2564A7FC60F3C9.taxon	biology_ecology	Movements, Home range and Social organization. Golden Langurs live mostly in unimale-multifemale groups that average 12 - 5 individuals, but vary in size from three to 40. Adult male-to-female sex ratios are 1: 1 - 5 - 3 - 3. Females are philopatric, but males disperse and are either solitary or form all-male groups before taking over as the dominant male in a new group. There is no evidentterritorial behavior, ranges overlap, and intergroup relations are relatively relaxed. Golden Langurs may even intermingle and travel together. Home ranges are relatively large at 150 - 600 ha. Population densities of Golden Langurs vary from 8 ind / km? in Manas National Park to 46 ind / km? in Ripu and 64 ind / km? ® in Chirrang.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB0FFB5FF2564A7FC60F3C9.taxon	conservation	Status and Conservation. CITES Appendix I. Classified as Endangered on The IUCN Red List. In India, the Golden Langur is included in Schedule I, Part I of the Indian Wildlife (Protection) Act, 1972, amended up to 2002. The Golden Langur is occasionally hunted but threatened mainly by loss of habitat. In 2006, the population was estimated at less than 1500 individuals in India and ¢. 4000 in Bhutan, with less than 2500 mature individuals in all. The Golden Langur occurs in Black Mountains, Royal Manas, and Thrumshingla national parks and Phipsoo Wildlife Sanctuary in Bhutan and Chakrashila Wildlife Sanctuary, Manas National Park, and Chirrang and Ripu forest reserves in India.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB0FFB6FF23696EFBC0F32F.taxon	materials_examined	type received from Barrackpore (menagerie), stated to be Malayan. Restricted by C. P. Groves in 2005 to India, Assam.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB0FFB6FF23696EFBC0F32F.taxon	discussion	T. pileatus forms, together with T. shortridgei and T: geei, the pileatus species group. The phylogenetic position of this group remains unresolved, and the species might have evolved through hybridization of ancestors (3 - 4 = 7 - 1 million years ago) of today’s genera Trachypithecus and Semnopithecus. It is thought that seasonal variation in fur color may accountfor at least some of the subspecies recognized. More fieldwork is needed to verify their validity. The subspecies tenebricus hybridizes with 7. gee: in Bhutan. Four subspecies recognized.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB0FFB6FF23696EFBC0F32F.taxon	distribution	Subspecies and Distribution. 1. p. pileatus Blyth, 1843 — NE India highlands S and E of the Brahmaputra River, in the states of Arunachal Pradesh, Assam, Meghalaya, and Nagaland (Karbi Anglong Plateau, Barail Range, and Khasi, Garo, Naga, and Jaintia Hills), and in NW Myanmar (W of the Chindwin River, S to Chin Hills Mts and Mt Victoria); the elevational range is 600 - 3000 m. 1. p. brahma Wroughton, 1916 — NE India, known only from the Dafla Hills, N of the Brahmaputra River, in Arunachal Pradesh State. 1: p. durga Wroughton, 1916 — E Bangladesh and NE India in the states of Assam, Mizoram, and Tripura (Naga Hills, Lakhimpur, Golaghat, Cachar Hills, Samaguting, and Sibsagar), adjoining the distribution of 7. p. pileatus to the N, but at lower elevations (i. e. from nearly sea level up to 600 m). 1. p. tenebricus Hinton, 1923 — NE India (Assam State) and Bhutan, in the Manas region N of the Brahmaputra River, with an elevational range of 100 - 2000 m.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB0FFB6FF23696EFBC0F32F.taxon	description	Descriptive notes. Head-body 53 - 72 cm (males) and 47 - 66 cm (females), tail 83 — 104 cm (males) and 72 - 102 cm (females); weight 11.5 - 14 kg (males) and 9.5 - 11.5 kg (females). The Capped Languris a large and highly variable form, noted forits short, upright crown hair. Coat is contrastingly colored, with a gray-to-brown dorsum and white-to-red underparts (belly often redder than chest and throat). Long, backwardswept cheek whiskers are buffy-white to red; crown cap is normally dark. External scrotum is absent. The “ Blond-bellied Capped Langur ” (1. p. pileatus) is iron-gray to pale brown above, with a yellowy-gray-white belly. Its chest and throat are whitish tinged with buff, and cheek whiskers are buffy-whitish, sharply contrasting with dark tone of dorsum. Its upper arms and entire legs are lighter, and hands, feet, and tail-tip are dark. Its crown cap is like the dorsum. The “ Orange-bellied Capped Langur ” (7. p. durga) 1 s pale slaty-gray above, with underparts (including inner surfaces of limbs), throat, and cheek whiskers orange to gingery, this tone extending further up flanks than in the Blond-bellied Capped Langur. Its head is not dark. The real existence of this subspecies is in question; its color tone perhaps merely the result of variations in temperature or humidity. The “ Buff-bellied Capped Langur ” (7. p. brahma) is less contrastingly colored than other subspecies, being generally dark gray above and becoming fawn-gray on lower flanks, rump, and legs. Its belly is white tinged with buff, chest and throat are white, and insides of limbs are pale gray. Its tail, hands, and feet are black. Its crown is not dark. The “ Tenebrous Capped Langur ” (7. p. tenebricus) is smaller and darker than the other subspecies, being generally ashy-black above, with a gray collar and legs. Belly is whitish in males and pale red in females, with remainder of underparts yellowish-orange. Its whiskers are grayish-yellow, and its black crown cap is defined behind a gray collar.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB0FFB6FF23696EFBC0F32F.taxon	biology_ecology	Habitat. Subtropical, broadleaf, evergreen, deciduous, submontane and bamboo forests and teak plantations. Depending on the region, the Capped Langur is sympatric with Phayre’s Langur (71. phayrei) and the Nepal Sacred Langur (Semnopithecus schistaceus).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB0FFB6FF23696EFBC0F32F.taxon	food_feeding	Food and Feeding. The Capped Langur is largely folivorous and eats young and mature leaves, supplemented with fruits, bark, seeds, flowers, and caterpillars. Gums and termite soil trails are also eaten on occasion. Water is obtained from tree cavities. The composition of the diet varies between seasons. Diets of the Orange-bellied Capped Langur have been studied at Madhupur in north-central Bangladesh. Leaves and leaf parts constituted 53 % of their overall diet (based on feeding records), followed by seeds and fruit (34 %), flowers (7 %), and other items such as buds and bark (6 %) and one caterpillar. Mature leaves accounted for 42 % of the diet and young leaves 11 %. Over a 15 month study, 35 species of trees, lianas, and vines were identified as food sources. Twenty-two species provided leaves, eleven fruits, seven flowers, and six seeds. The mature leaf component of the diet varied from 5 - 1 % in March-April when new leaves and fruits were abundant at the beginning of the monsoon to 65 - 1 % in late winter in January-February when about two-thirds of the trees had lost their leaves. In May-September, fruits accounted for nearly 50 % of the diet. Important species providing fruits were Ficus benghalensis and F. religiosa (Moraceae), Spondias mangifera (Anacardiaceae), and Terminalia bellirica (Combretaceae). Two species provided seeds, Mallotus philippensis (Euphorbiaceae) and Litsea polyantha (Lauraceae). At Pakhui Wildlife Sanctuary, the Tenebrous Capped Langur spent 67 % ofits annual feeding time eating leaves: 57 % young leaves, 6 % mature leaves, and 4 % petioles. They also ate fruits and seeds (16 %) and flowers and buds (16 %). Fifty-two plant species in 30 families were included in their diets. The most important species during the year were Gmelina arborea (Verbenacae; leaves), Albizia lucida (Fabaceae; leaves and fruits), Ficus glomerata (Moraceae; leaves), Mikania micrantha (Asteraceae; leaves), and Morus laevigata (Moraceae; leaves). Together, they accounted for nearly 60 % of the feeding records. Most of their feeding (68 %) was in the upper canopy. At Pakhui, they also fed, for some hours at a time, on water lilies.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB0FFB6FF23696EFBC0F32F.taxon	breeding	Breeding. At Pakhui Wildlife Sanctuary in Arunachal Pradesh, the Tenebrous Capped Langur has two breeding seasons: September — January and April-May. At Madhupur, breeding occurs in October — February and later in April. At both sites, breeding occurred in the dry season. Females show proceptive behavior by head shaking, presenting their hindquarters to males, and raising an arm in a distinct fashion. This behavior sometimes leads to aggression, and males are aggressive to females or chase them before mating with them. Females give birth to a single young from late December through April; ¢. 50 % of births occur in February-March. The gestation period averages 200 days (196 - 205 days). The interbirth interval is c. 2 years. Infants up to five months of age are creamy-white, with a golden tinge and pink skin. Allomothering takes place among adult females but is restricted to parous females; nulliparous females are rarely allowed to take infants. Allomothering is also restricted to particular female dyads; newborn infants are transferred to another female, but a single one during the first three months. This behavior is seen as altruistic, allowing lactating females to feed.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB0FFB6FF23696EFBC0F32F.taxon	activity	Activity patterns. Capped Langurs are diurnal and mainly arboreal. They occasionally feed and play on the ground. They feed early (06: 00 - 10: 00 h) and late (15: 00 - 19: 00 h) in the day, and rest at midday (11: 00 - 15: 00 h). Resting takes up 40 % of their activity budget, feeding 35 %, traveling 18 %, and playing, grooming, and engaging in other activities 6: 8 %. Time spent feeding changes seasonally and is dependent on the contribution of fruit in the diet (which can be harvested more rapidly). In some months, feeding can be as low as 26 % oftheir daily activities, and in others as high as 41 %.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB0FFB6FF23696EFBC0F32F.taxon	biology_ecology	Movements, Home range and Social organization. Capped Langurs live in unimale — multifemale groups, with an average 7 - 5 individuals (range 2 - 14). All-male groups and solitary males are also observed. Home ranges vary among localities (c. 21 - 64 ha). Daily movements are 320 - 800 m. Capped Langurs are territorial, and the adult male dedicates much of his day to vigilance. While a group rests, the male goes up into the canopy and calls and spreads his legs on either side of a branch to expose his lighter bare skin and give a penile display. Groups within 50 m of each other cause tension, and a male may chase his own group members away. Males display to each other by moving rapidly through trees and emitting a “ hak ” call. Agitated females scream and hug each other. A female group member approaching an opposing group might be chased and attacked by her group’s male; he will slap, bite, and pull at her if he catches her. Sometimes females transfer to other groups. Solitary males follow bisexual groups and try to gain access to the females. The resident male is less aggressive to males with whom he is familiar; he responds to the approach of unfamiliar males with more displaying than he does with familiar males. The pattern of male competition appears to involve successfully keeping females in the group, rather than aggressive confrontation with males. Golden Jackals (Canis aureus) will attack Capped Langurs when they are on the ground.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB0FFB6FF23696EFBC0F32F.taxon	conservation	Status and Conservation. CITES Appendix I. Classified as Vulnerable on The IUCN Red List. The subspecies pileatus, durga, and tenebricus are classified as Endangered, and brahma as Data Deficient. Capped Langurs are nationally protected in all the countries where they occur. Itis the most common langur in north-eastern India, but habitat loss and hunting are major threats throughout its range. Habitat loss and fragmentation are caused mainly by expanding agriculture and timber and firewood collection; however, Capped Langurs are more adaptable to this situation than the sympatric Western Hoolock Gibbon (Hoolock hoolock). They are captured for the pet trade and widely hunted for their meat and body parts. The Orange-bellied Capped Langur is found in Madhupur National Park and Chunati, Kalenga and Rema wildlife sanctuaries in Bangladesh and Dampa Tiger Reserve, Murlen and Kaziranga national parks and Sepahijala, Trishna, Gumti Wildlife, Nengpui, Pabitora and Gibbon wildlife sanctuaries in India. The Tenebrous Capped Langur is found in Royal Manas National Park in Bhutan and Manas and Nameri national parks and Eagle Nest and Pakhui wildlife sanctuaries in India. The Blond-bellied Capped Langur is found in Namdapha, Balphakram, and Nokrek national parks and Siju Wildlife Sanctuary in India. A population estimate for the Capped Langur suggests less than 5600 mature individuals. For Blond-bellied and Orange-bellied capped langurs, fewer than 2500 mature individuals are estimated, with subpopulations having less than 250 mature individuals. The population of Tenebrous Capped Langurs is thought to be less than 600 mature individuals, and the Buff-bellied Capped Langur is only known from the holotype that was collected in 1911. Survey work to determine the population and conservation status is needed, particularly for the Buff-bellied Capped Langur.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB3FFB6FFE9680BF6E4F7A7.taxon	materials_examined	Burma, Homalin, upper Chindwin.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB3FFB6FFE9680BF6E4F7A7.taxon	discussion	T. shortridgei forms, together with T. pileatus and T. geei, the pileatus species group. The phylogenetic position of this group remains unresolved, and they might have evolved through hybridization of ancestors (3 - 4 - 7 - 1 million years ago) of today’s genera Trachypithecus and Semnopithecus. In the Chindwin headwaters, there might pe intergradation with 7. pileatus or an undescribed variation of 1. shortridgei. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB3FFB6FFE9680BF6E4F7A7.taxon	distribution	Distribution. NE Myanmar, E of the Chindwin River (Kachin State N to Myitkyina District) and SW China (Dulong River Valley in Gongshan County, NW Yunnan Province).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB3FFB6FFE9680BF6E4F7A7.taxon	description	Descriptive notes. Head — body 67 - 72 cm, tail 95 - 104 cm; no specific data are available for body weight. Shortridge’s Langur is strikingly uniform in color, being silvery-gray on head and body (including underside, crown, and cheek whiskers), with lighter legs. Tail becomes increasingly dark toward tip. Hands and feet are black. Backward-swept cheek whiskers are shorter than those of Capped Langurs (71. pileatus) and Golden Langurs (71. geet), and they contrast with short, upright crown hair. External scrotum is absent.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB3FFB6FFE9680BF6E4F7A7.taxon	biology_ecology	Habitat. Lowland evergreen and semi-evergreen forest at elevations of 200 - 2500 m. Shortridge’s Languris rarely encountered at higher elevations. It is sympatric with the Stryker’s Snub-nosed Monkey (Rhinopithecus strykerr).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB3FFB6FFE9680BF6E4F7A7.taxon	food_feeding	Food and Feeding. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB3FFB6FFE9680BF6E4F7A7.taxon	breeding	Breeding. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB3FFB6FFE9680BF6E4F7A7.taxon	activity	Activity patterns. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB3FFB6FFE9680BF6E4F7A7.taxon	biology_ecology	Movements, Home range and Social organization. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB3FFB6FFE9680BF6E4F7A7.taxon	conservation	Status and Conservation. CITES Appendix I. Classified as Endangered on The [UCN Red List. Shortridge’s Languris protected in Myanmar and China and occurs in at least three protected areas: Hponkanrazi Wildlife Sanctuary and Hkakaborazi National Park in Myanmar and Gaoligongshan National Nature Reserve in China. Langurs occur also in Hukaung Tiger Reserve, but the species is not known. Population numbers of Shortridge’s Langur are unknown, but they are believed to be declining because of habitat loss and hunting. They are hunted for meat and use in traditional medicine, and they are sometimes exported illegally to China. Habitat loss from agriculture, logging, and firewood extraction and the planned Tazone hydroelectric power dam at the Chindwin River are major threats. The latter will result in direct loss of a large part of the remaining habitat used by Shortridge’s Langur and will harm populations via an increase of hunting by dam construction workers. If Shortridge’s Langur does not occur in Hukaung Tiger Reserve, which provides a large area of suitable lowland habitat, it is seriously threatened because most other protected areas in Myanmar are highland habitats, and suitable habitat below 900 m is heavily degraded and fragmented. There is an urgent need for further survey work to determine current population status, limits of distribution, and habitat preferences of this poorly known species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB3FFB7FAF06DB4FE5FFB0E.taxon	materials_examined	Indonesia, Semarang. Rrestricted by D. Brandon-Jones in 1995 to Batu District.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB3FFB7FAF06DB4FE5FFB0E.taxon	discussion	Until recently, 7. mauritius was considered to be a subspecies of 1. auratus, but genetic data support a separation at the species level. T. auratus is closely related to T. cnistatus, 1. germaini, 1. margarita, 1. mauritius, and T. selangorensis, and together they form the cristatus speciesgroup. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB3FFB7FAF06DB4FE5FFB0E.taxon	distribution	Distribution. E Java (NW to the vicinity ofJakarta, and E of ¢. 109 ° E in the S coast), Bali, and Lombok Is, and Pulau Sempu and Nusa Barung Is, off the S coast ofJava.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB3FFB7FAF06DB4FE5FFB0E.taxon	description	Descriptive notes. Head-body 44 - 65 cm (males), tail 61 - 87 cm (males); weight c. 7 - 1 kg (males); no measurements are available for females. The EastJavan Langur is a large and extraordinarily beautiful monkey. Its circumfacial hair is erect and forwardcurled, facial skin is bluish or blackish, and ears have whitish or yellowish tufts. Females differ from males in having a pale, usually yellowish-white pubic patch. Jaws and teeth are notably large. There are two main color morphs; the more common is glossy-black, with a slight brownish tinge (particularly on belly, sideburns, and legs). Its hairs are light-tipped on arms, legs, head, flanks, and sometimes back. Palms, soles, and facial skin are slaty-black. The much rarer “ red ” morph is restricted to an area between Blitar, [jen, and Pugeran. It is deep orange, becoming more yellowish on limbs, ears, and belly. There is often a black fringe on its dorsum, and its palms, soles, and facial skin are depigmented or freckled.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB3FFB7FAF06DB4FE5FFB0E.taxon	biology_ecology	Habitat. Remnant primary and secondary, dry deciduous, mangrove, beach, freshwater swamp, and hill forests at elevations up to 3500 m. The East Javan Langur is sometimes found in teak, rasamala, and Acacia plantations. It is sympatric with the “ Javan Fuscous Langur ” (Presbytis comata fredericae).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB3FFB7FAF06DB4FE5FFB0E.taxon	food_feeding	Food and Feeding. The EastJavan Langur is mostly folivorous, feeding on young leaves (46 %), ripe fruits (27 %), unripe fruits (8 %), flowers (7 %), mature leaves (1 %), and animal prey (1 %, insects and larvae). In areas where native vegetation has been replaced by teak plantations, it has learned to feed on teak leaves.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB3FFB7FAF06DB4FE5FFB0E.taxon	breeding	Breeding. EastJavan Langurs mate throughoutthe year. A single young is born. Infants are entirely orange, with pale skin on their faces and extremities. Allomothering is common.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB3FFB7FAF06DB4FE5FFB0E.taxon	activity	Activity patterns. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB3FFB7FAF06DB4FE5FFB0E.taxon	biology_ecology	Movements, Home range and Social organization. Home range sizes of East Javan Langurs differ among locations and vary from 2 - 5 - 8 ha to 20 - 30 ha, with little overlap. Group sizes are 5 - 23 individuals, with an average of 7 - 9 individuals. EastJavan Langurs live mainly in unimale-multifemale groups, but two males have been observed in mixed groups. Males sometimes form all-male groups.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB3FFB7FAF06DB4FE5FFB0E.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Vulnerable on The IUCN Red List (as 1. auratus auratus). The EastJavan Langur has been protected by Indonesian law since 1999. Although still widely distributed, populations are reduced and fragmented. General habitat loss and degradation due to agriculture and human settlements are major threats, as are hunting for food and the pet trade. It occurs in at least seven protected areas in Indonesia: Alas Purwo, Bali Barat, Baluran, BromoTengger Semeru, Gunung Merapi, and Meru Betiri national parks and Pangandaran Nature Reserve.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB2FFB7FF226941F8C0F646.taxon	materials_examined	Indonesia, Sumatra, Bencoolen (= Bengkulu).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB2FFB7FF226941F8C0F646.taxon	discussion	T. selangorensis was first described as a subspecies of T. cristatus, but is now considered a distinct species. T. cristatus is closely related to T. auratus, T. mauritius, T. germaini, 1. margarita, and T. selangorensis, and together they form the cristatus species group. Two subspecies are recognized, although they are genetically very similar.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB2FFB7FF226941F8C0F646.taxon	distribution	Subspecies and Distribution. 1. c. cnistatus Raffles, 1821 — Borneo, Sumatra, Bangka, Belitung, and Riau (Bintan, Combol, Sugi, and perhaps Batam) and Lingga (Bakung, Lingga) Archipelagos, off E Sumatra. 1: c. vigilans G. S. Miller, 1913 — Natuna Is (Serasan I), off W Borneo.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB2FFB7FF226941F8C0F646.taxon	description	Descriptive notes. Head — body 50 - 58 cm (males) and 46 - 51 cm (females), tail 67 — 75 cm (males) and 66 - 75 cm (females); weight ¢. 6 - 6 kg (males) and ¢. 5 - 7 kg (females). The Silvered Languris variable in color. Circumfacial hair is erect and forward-curled, there is a high pointed crest (but less pointed and high that in the Selangor Silvery Langur, 1. selangorensis), and face is slaty-black. Adult female Silvered Langurs have irregular patches of white on inner parts of upper thighs. Teeth and jaws are relatively short. The nominate subspecies cristatus can be divided into two main morphs, which are very distinct but not genetically and geographically separated (although they are somewhat more so in Sumatra). The silvery-gray “ wltimus ” morph predominates in northern Sumatra; it is uniformly dark gray, tipped with paler gray; it has a black-tinged forehead and yellowish groin and underside oftail. The “ cristatus ” morph has shorter hair tips, especially on its back, giving it an overall almost gray-green color. It occurs further south on Sumatra and on Bangka, Belitung, and Riau Islands. Both morphs occur at almost equal frequencies in Borneo, with the “ cristatus ” morph being perhaps slightly more common. There is an additional, rare reddish morph in Sabah, Borneo. “ Miller’s Silvered Langur ” (7. c. vigilans) is rather darker than the nominate subspecies, with less lightened hair tips. Its crown is dark and surrounded by a well-defined zone of long hair tips.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB2FFB7FF226941F8C0F646.taxon	biology_ecology	Habitat. Primary and secondary forests including coastal, montane, mangrove, swamp, and riparian forest in Sumatra and mainly riparian and mangrove forests in Borneo. The Silvered Langur is sometimes found in plantations. Depending on the region, it is sympatric with different species of Presbytis.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB2FFB7FF226941F8C0F646.taxon	food_feeding	Food and Feeding. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB2FFB7FF226941F8C0F646.taxon	breeding	Breeding. The Silvered Langur is not a seasonal breeder. The gestation period averages 194 - 6 days, and the ovarian cycle lasts c. 24 days. It usually has singletons (twins are rare). Newborns are golden-orange in color, with white on the hands, feet, and face. [ts coat gradually changes to grayish at 3 - 5 months of age. Allomothering is common.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB2FFB7FF226941F8C0F646.taxon	activity	Activity patterns. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB2FFB7FF226941F8C0F646.taxon	biology_ecology	Movements, Home range and Social organization. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB2FFB7FF226941F8C0F646.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Near Threatened on The IUCN Red List, including the nominate subspecies cristatus, but the subspecies vigilans is classified as Critically Endangered. Major risks for Miller’s Silvered Langur are its restricted distribution (only 62 km * on Serasan Island) and ongoing loss of suitable habitat due to deforestation. The nominate subspeciesis still wide-ranging, but it too is suffering from habitat loss throughoutits range. On Borneo, threats are mainly due to land clearance (oil palm plantations), and forest fires. Silvered Langurs are hunted for their meat in Sarawak and are often kept as pets in Sumatra. The nominate subspecies occurs in various protected areas, including Bukit Barisan Selatan and Gunung Leuser national parks in Indonesia and Bako National Park in Malaysia. An overall population estimate is not available.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB2FFB7FF25602CFEF8F3FB.taxon	materials_examined	“ Mauritius, ” probably W Java. Restricted by D. Brandon-Jones in 1995 to Jasinga.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB2FFB7FF25602CFEF8F3FB.taxon	discussion	Until recently, 7 " mauritius was considered a subspecies of 1. auratus, but genetic data support a separation at species level. 1 " mauritius is closely related to T. cristatus, 1. auratus, 1. germaini, 1. margarita, and 1. selangorensis, and together they form the eristatus species group. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB2FFB7FF25602CFEF8F3FB.taxon	distribution	Distribution. W Java (restricted to the N coast W ofJakarta, and inland to Bogor, Cisalak, andJasinga, SW to Ujung Kulon, then along the S coastas far E as Cikaso and possibly Ciwangi).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB2FFB7FF25602CFEF8F3FB.taxon	description	Descriptive notes. There are no specific measurements available, but the West Javan Langur is probably similar to the East Javan Langur (7 auratus). Circumfacial hair of the West Javan Langur is erect and forward-curled, facial skin is bluish or blackish, and ears have whitish or yellowish tufts. Females differ from males in having a pale, usually yellowish-white, pubic patch. Jaws and teeth are notably large. Palms and soles are slaty-black. Fur is glossy-black, with a very slight brownish tinge, especially on belly, sideburns, and legs. The West Javan Langur is generally similar to the “ black ” morph of the East Javan Langur, but it lacks the light-tipped hairs.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB2FFB7FF25602CFEF8F3FB.taxon	biology_ecology	Habitat. Remnant primary and secondary dry deciduous, mangrove, beach, freshwater swamp, and hill forests. The West Javan Langur is sympatric with the Javan Langur (Presbytis comata).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB2FFB7FF25602CFEF8F3FB.taxon	food_feeding	Food and Feeding. There is no specific information available for this species, but the diet of the WestJavan Langur is presumably similar to that of the East Javan Langur.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB2FFB7FF25602CFEF8F3FB.taxon	breeding	Breeding. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB2FFB7FF25602CFEF8F3FB.taxon	activity	Activity patterns. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB2FFB7FF25602CFEF8F3FB.taxon	biology_ecology	Movements, Home range and Social organization. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB2FFB7FF25602CFEF8F3FB.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Vulnerable on The IUCN Red List (as 1. auratus mauritius). The West Javan Langur has been protected by Indonesian law since 1999. Although still widely distributed, its forest habitats are much reduced and fragmented due to agriculture and human settlements. The West Javan Langur is hunted for food and the pet trade. It occurs in Gunung Gede Pangrango, Gunung Halimun Salak, and Ujung Kulon national parks, Cibanteng Nature Reserve, and Cikepuh Game Reserve.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB2FFB0FA2F6F14FDBAFBA6.taxon	materials_examined	West Malaysia, Selangor Province, Kuala Selangor (3 ° 20 ° N, 101 ° 20 ’ E).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB2FFB0FA2F6F14FDBAFBA6.taxon	discussion	T. selangorensis is closely related to T. cristatus, T. auratus, T. mauritius, T. germaini, and T. margarita, and together they form the cristatus species group. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB2FFB0FA2F6F14FDBAFBA6.taxon	distribution	Distribution. W Peninsular Malaysia, confined to a Strip along the W coast from 6 ° N to 1 ° 30 ° N in the states of Kedah, Perak, Selangor, Negeri Sembilan, Melaka, and Johor.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB2FFB0FA2F6F14FDBAFBA6.taxon	description	Descriptive notes. Head-body 50.6 - 53.5 cm (males) and 45 - 8 cm (females), tail 70.4 - 82.3 cm (males) and 68 cm (females); no specific data are available for body weight. Body of the Selangor Silvery Langur is a uniform dark gray, and facial skin, hands, and feet are black. Face has no clear eye-rings. Gray whiskers are long and straight, and thus, clearly differ from the mussel-shaped whiskers in the Silvered Langur (1. cristatus).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB2FFB0FA2F6F14FDBAFBA6.taxon	biology_ecology	Habitat. Mainly riparian and mangrove forest, and occasionally plantations. The Selangor Silvery Langur is sympatric with the Dusky Langur (7. obscurus), the Banded Langur (Presbytis femoralis), and the Pale-thighed Langur (P. siamenss).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB2FFB0FA2F6F14FDBAFBA6.taxon	food_feeding	Food and Feeding. The Selangor Silvery Langur is mainly folivorous, with leaves comprising up to 91 % of its diet. It also eats fruits, seeds, flowers, and young shoots.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB2FFB0FA2F6F14FDBAFBA6.taxon	breeding	Breeding. There is no specific information available for this species, but the Selangor Silvery Langur is not a seasonal breeder. Neonatal Selangor Silvery Langurs are born with orange fur, and white skin on hands, feet, and face.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB2FFB0FA2F6F14FDBAFBA6.taxon	activity	Activity patterns. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB2FFB0FA2F6F14FDBAFBA6.taxon	biology_ecology	Movements, Home range and Social organization. The Selangor Silvery Langur lives in unimale-multifemale groups of 11 - 38 individuals (average 26 - 5), and smaller allmale groups. Home ranges are 20 - 43 ha. Adjacent home ranges overlap, but they are defended. Selangor Silvery Langurs travel only 200 - 500 m / day, usually led by a male. Lone males have also been observed. When a new male enters a group, infanticide might occur although it has not been confirmed.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB2FFB0FA2F6F14FDBAFBA6.taxon	conservation	Status and Conservation. CITES Appendix II. The Selangor Silvery Langur has not been assessed on The IUCN Red List. A major threat is habitat loss due to land clearance and forest fires, specifically in coastal regions. It occurs in Kuala Selangor and Bukit Nanas nature reserves in Peninsular Malaysia.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB5FFB0FFF261B3F791FD0B.taxon	materials_examined	Cochin China and Cambodia.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB5FFB0FFF261B3F791FD0B.taxon	discussion	Until recently, T. margarita was considered a synonym of T. germaini. T. germaini is closely related to T. cristatus, T. auratus, T. mauritius, T. margarita, and T. selangorensis, and together they form the cristatus species group. It is partially sympatric with T. barbei. There is a highly distinct but little-known taxon from northern Vietnam that may be attributable to this species (as the form caudalis). Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB5FFB0FFF261B3F791FD0B.taxon	distribution	Distribution. Mainland South-east Asia in S Myanmar, S Thailand, S Laos, Cambodia (W of Mekong River), and the S tip of Vietnam. Range limit of this species to the E might be the Mekong River.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB5FFB0FFF261B3F791FD0B.taxon	description	Descriptive notes. Head — body ¢. 55 cm, tail 72 - 84 cm; weight 6.5 - 7 kg. Germain’s Langur is similar to the Silvered Langur (7. ¢ ristatus), but larger and paler and has a longer tail. Fur is generally pale grayish, with a lighter underside, throat, and shanks. Forearms are a dark gray, hands and feet are black, and tail is black above and lighter below. Long, creamy-yellow circumfacial hairs form a distinctive “ halo. ” Face is slaty-black. Females have a whitish pubic patch. The little-known form from northern Vietnam is described as dark, with a black tone to its dorsum, becoming brown-gray with silvery tips on flanks. Long cheek hairs are pale yellow, and legs are pale yellowish on their outsides. Its tail is bi-colored at the base, with a sharply marked yellowish stripe underneath, contrasting with the black upper side (although not distally black).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB5FFB0FFF261B3F791FD0B.taxon	biology_ecology	Habitat. Evergreen, semi-evergreen, mixed deciduous, riparian, and gallery forest. Records of Germain’s Langurs in hilly areas or at higher elevations are few. Depending on the region, it is sympatric with the Tenasserim Langur (71. barber).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB5FFB0FFF261B3F791FD0B.taxon	food_feeding	Food and Feeding. Germain’s Langurs eat mainly young leaves, along with shoots, seeds, flowers, dried wood, crops, and fruits (especially figs). Soil is also consumed occasionally. It has not been observed eating animal prey.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB5FFB0FFF261B3F791FD0B.taxon	breeding	Breeding. Infant Germain’s Langurs have a golden-orange coat, with white faces, hands, and feet.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB5FFB0FFF261B3F791FD0B.taxon	activity	Activity patterns. Germain’s Langur is diurnal and arboreal.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB5FFB0FFF261B3F791FD0B.taxon	biology_ecology	Movements, Home range and Social organization. Germain’s Langurs live in groups of 9 - 30 individuals (average 17). Aspects of social organization are likely similar to other members of the cristatus species group.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB5FFB0FFF261B3F791FD0B.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Endangered on The IUCN Red List. Germain’s Langur is widespread but relatively rare throughoutits range. Population densities are generally low. In Cambodia, it is widespread in certain areas but rare in others. In Thailand, it is moderately common in several protected areas but has declined significantly in recent years. In Vietnam, five or six karst hills in Kien Giang Province hold the only confirmed populations of Germain’s Langurs in the country, and excavation of limestone for cement production in this area is a serious threat. The major threats throughout its range are hunting (food, traditional medicine, and the pet trade) and habitat loss due to agriculture. It occurs in several protected areas in Thailand and Cambodia. In Laos, it is confirmed only in Dong Khanthung National Biodiversity Conservation Area. Population estimates are not available. Additional surveys and taxonomic work are needed to elucidate the population and conservation status and distributional limits of Germain’s Langur.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB5FFB0FAFF662FF8E9F46A.taxon	materials_examined	Vietnam, Langbian.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB5FFB0FAFF662FF8E9F46A.taxon	discussion	Until recently, T. margarita was considered a synonym of T. germaini, but it was elevated to a full species based on genetic data. T. margarita is closely related to T. cristatus, T. auratus, T. mauritius, T. germaini, and T. selangorensis, and together they form the cristatus species group. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB5FFB0FAFF662FF8E9F46A.taxon	distribution	Distribution. Mainland South-east Asia in S Laos (N limit is 16 ° 23 ’ N), SC Vietnam (N limit is 14 ° 30 ’ N), and E Cambodia (Ratanakiri and Mondulkiri provinces); most likely the W limitis the Mekong River, but further studies are needed to confirm this. Records in Vietnam N to 16 ° 37 ° N are questionable.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB5FFB0FAFF662FF8E9F46A.taxon	description	Descriptive notes. There are no specific measurements available, but the Annamese Languris similar in size to Germain’s Langur (7. germaini). Fur of the Annamese Langur is generally pale grayish, with underside and throat very light, contrasting with black forehead, forearms, hands, and feet. Face is gray, with white eye-rings in many individuals. Females have a whitish pubic patch.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB5FFB0FAFF662FF8E9F46A.taxon	biology_ecology	Habitat. Lowland evergreen, semi-evergreen, mixed deciduous, riparian, and gallery forest. Records of the Annamese Langur in hilly areas or at higher elevations are few. It is sympatric with doucs (Pygathrix).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB5FFB0FAFF662FF8E9F46A.taxon	food_feeding	Food and Feeding. There is no specific information available for this species, but the Annamese Langur is probably primarily folivorous, as are other members of the cristatus species group.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB5FFB0FAFF662FF8E9F46A.taxon	breeding	Breeding. Infant Annamese Langurs have a golden-orange coat, with a white face, hands, and feet.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB5FFB0FAFF662FF8E9F46A.taxon	activity	Activity patterns. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB5FFB0FAFF662FF8E9F46A.taxon	biology_ecology	Movements, Home range and Social organization. There is no specific information available for this species, but the Annamese Langur is most likely similar to other members of the cristatus species group.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB5FFB0FAFF662FF8E9F46A.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Endangered on The IUCN Red List (under 1. germaini). The Annamese Langur is widespread but rare throughout its range. Densities are generally low. Only a few sightings have been recorded in the last few decades in Vietnam, and it might be the rarest and most threatened monkey in Laos. In Cambodia, it is widespread in certain areas (e. g. Mondulkiri Province), but rare in others. Throughout its distribution, major threats are habitat loss due to agricultural land use and hunting for food, traditional medicine, and the pet trade. It occurs in numerous protected areas, among them: Dong Phou Vieng, Xe Bang Nouan, Xe Pian, and Dong Hua Sao national biodiversity conservation areas and Bolaven Southwest Proposed National Protected Area in Laos; Cat Tien, Yok Don, Chu Mom Ray, Chu Prong, and Kon Ka Kinh national parks and Takou Nature Reserve in Vietnam; and Virachey National Park, Mondulkiri Protected Forest, and Lomphat, Phom Prich, Phom Nam Lyr, and Snoul wildlife sanctuaries in Cambodia. Population estimates for Annamese Langurs are not available. Additional surveys and taxonomic work are needed to elucidate its population status and distributional limits.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB6FFB3FF30640AFC6BF88C.taxon	materials_examined	Burma, Ye, south of Moulmein.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB6FFB3FF30640AFC6BF88C.taxon	discussion	The rather mysterious forms originally known, respectively, as Presbytis argenteus and Semnopithecus holotephreus are variously treated under this species or with 7. phayre, and may be in some way distinct. The argenteus morph is silver, while holotephreus 1 s ashy-gray. 1. barbeiis a member of the obscurus species group, which also includes 1. obscurus, 1. crepusculus, and 1. phayre:. 1: phayrei occurs to the north, 71. obscurus to the south, and 7. germaini to the south-east, with which it is sympatric. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB6FFB3FF30640AFC6BF88C.taxon	distribution	Distribution. E Myanmar and W Thailand, confined to a small region N of the Thai-Malai Peninsula (14 ° 20 ° - 15 ° 10 ° N, 98 ° 30 ° - 98 ° 55 ’ E).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB6FFB3FF30640AFC6BF88C.taxon	description	Descriptive notes. There are no specific measurements available. General color of the Tenasserim Languris grayish black with no silvering, and only slightly lighter ventrally. Tail is dark gray, slightly paler than body. Root of tail and area around ischial callosities are whitish. Long, upright crown hairs form a distinct crest. Face is gray with a violet tinge. Whitish eye-rings fully encircle eyes, and there is depigmentation around mouth.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB6FFB3FF30640AFC6BF88C.taxon	biology_ecology	Habitat. Precise habitat preferences are unclear. Habitats within the distribution of the Tenasserim Langur include mixed deciduous dipterocarp and moist evergreen forest.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB6FFB3FF30640AFC6BF88C.taxon	food_feeding	Food and Feeding. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB6FFB3FF30640AFC6BF88C.taxon	breeding	Breeding. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB6FFB3FF30640AFC6BF88C.taxon	activity	Activity patterns. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB6FFB3FF30640AFC6BF88C.taxon	biology_ecology	Movements, Home range and Social organization. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB6FFB3FF30640AFC6BF88C.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Data Deficient on The [UCN Red List. The Tenasserim Langur is known only from museum specimens, some sightings in the wild dating back to the 1970 s, and three captive individuals. Its distribution is estimated to encompass 10,000 - 12,000 km? and it is not known to occur in any protected areas. Presumably hunting and habitat loss are major threats. There is an urgent need for additional surveys to determine its current population status, the limits of its distribution, and its habitat preferences.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB6FFB3FF3362A9FA5FF557.taxon	materials_examined	Malaysia. Restricted by F. N. Chasen in 1940 to Malacca.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB6FFB3FF3362A9FA5FF557.taxon	discussion	With T. phayrei, T. barbei, and T. crepusculus, I. obscurus forms the obscurus species group. The island subspecies, in particular, are quite similar to one another. Seven subspecies: recognized 2.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB6FFB3FF3362A9FA5FF557.taxon	distribution	Subspecies and Distribution. T. o. obscurusReid, 1837 — PeninsularMalaysia (SfromaboutPerlis). 1: o. carboThomas & Wroughton, 1909 — Langkawi, Terutau, andDayangBuntingIs, offtheWcoastoftheThai-MalayPeninsula. T: o. flavicauda Elliot, 1910 — S Myanmar (S of ¢. 15 ° 10 ’ N, and including James, Kisseraing, and King Is in the Mergui Archipelago), SW Thailand (including Khao Lak I), and N Peninsular Malaysia (S to Perlis). 1: o. halonifer Cantor, 1845 — Penang I, off W Peninsular Malaysia. 1: o. sanctorum Elliot, 1910 — Mergui Archipelago (Saint Matthew I), off W Myanmar. 1 o. sesemundi Chasen, 1940 — Ko Phangan I; it also may occur on the adjacent E coast of the Thai — - Malay Peninsula. I: o. styx Kloss, 1911 — East Perhentian I; it also may occur on the adjacent E coast of the Thai-Malay Peninsula.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB6FFB3FF3362A9FA5FF557.taxon	description	Descriptive notes. Head-body 42 - 68 cm (males) and 43 - 60 cm (females), tail 64 — 81 cm (males) and 57 - 79 cm (females); weight 6.1 - 9.1 kg (males) and 5.8 - 6 kg (females). Coloration of the Dusky Langur varies considerably, but it is generally grayishblue or black above, with a lighter underside and contrasting whitish or gray on legs and tail, which may also be yellow. Some subspecies can have a prominent light or white crown. Face is dark gray, with characteristic “ spectacles ” of white skin surrounding eyes, and muzzle is whitish or pink. Sexes are generally similar in appearance, although males are somewhat heavier. “ Reid’s Dusky Langur ” (7. o. obscurus) is bronze to medium drab-brown above, with limbs and tail contrastingly pale. There is a distinct dorsalstripe. Its posterior crown patch is creamy or yellow-brown. In the “ Blond-tailed Langur ” (1. o. flavicauda), there is considerable variation, with some individuals much less contrastingly patterned than others, although all are basically black with creamyyellow or brownish-gray on legs and tail. Adult females tend to be paler and browner than adult males, and subadult males are more blackish. “ Cantor’s Langur ” (1. o. halonifer) is generally brownish, with a distinct dorsal stripe. Its limbs and tail do not contrast greatly with its body. Its crown is a drab creamy-buff. The “ Perhentian Langur ” (7: o. styx) is uniformly deep brown, with no trace of a dorsal stripe. Its limbs and tail are only slightly lighter and grayer than its upper body. Crown is buffy. The “ Tarutao Langur ” (7. o. carbo) is similar to Cantor’s Langur but slightly darker, with its dorsal stripe less clearly marked. The “ Phangan Langur ” (7. o. seimundi) is a dark insular form similar to the Tarutao Langur but with a browner frontal band and slightly paler limbs. The “ Zadetkyi Kyun Langur ” (7. o. sanctorum) is black above, with a creamy colored posterior chest. Its arms are black, legs are dark smoky-gray, and feet are black. Its tail is yellow-gray.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB6FFB3FF3362A9FA5FF557.taxon	biology_ecology	Habitat. Primary, secondary, lowland, submontane, and montane forest, up to elevations of 1800 m. The Dusky Langur seems to prefer old growth forests. It can also be found in scrub areas, rubber tree plantations, and gardens. Depending on the region, it is sympatric with the Selangor Silvery Langur (7. selangorensis), the Banded Langur (Presbytis femoralis), and the Pale-thighed Langur (P. siamensis).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB6FFB3FF3362A9FA5FF557.taxon	food_feeding	Food and Feeding. The Dusky Langur is largely folivorous. Its diet contains leaves (58 %), fruits (mainly figs) and seeds (35 %), and flowers (7 %). Insects are occasionally eaten. Groups of Dusky Langurs frequently raid rubber plantations, where they are capable of causing significant damage in a short period of time. Individuals can eat upwards of two kilograms of vegetation daily. Groups split up into subgroups to feed.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB6FFB3FF3362A9FA5FF557.taxon	breeding	Breeding. The Dusky Langur most likely is not a seasonal breeder. Females do not show a sexual swelling during the periovulatory period of their ovarian cycle. A single young is born after a gestation of c ¢. 150 days. Infants are covered by a bright apricot natal coat that contrasts sharply with adult coloration. All adults participate in raising offspring, and allomothering has been observed.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB6FFB3FF3362A9FA5FF557.taxon	activity	Activity patterns. The Dusky Langur lives and feeds in the high canopy (25 - 50 m). It spends ¢. 46 % of its day resting, 34 % feeding, and 20 % traveling.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB6FFB3FF3362A9FA5FF557.taxon	biology_ecology	Movements, Home range and Social organization. The Dusky Langur lives in unimale — multifemale or multimale-multifemale groups of 10 - 17 individuals, with an average of 2 - 5 males / group. It moves ¢. 950 m / day and has a home range of c. 33 ha. All-male groups and solitary males also have been observed. Population densities on the mainland are estimated at 1 - 8 - 7 - 3 groups / km?. Locomotion consists of 50 % quadrupedal walking and running, 40 % leaping, and 9 % climbing.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB6FFB3FF3362A9FA5FF557.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Near Threatened on The IUCN Red List. The subspecies styx, seimundi, and sanctorum are classified as Vulnerable, and obscurus, flavicauda, halonifer, and carbo as Least Concern on The [UCN Red Lust. Major threats to Dusky Langurs are hunting for food and habitat loss and degradation, mainly because of expanding oil palm plantations, agriculture, urbanization, and touristic development. They are sometimes also killed in road accidents. For the insular forms, their restricted distributions make them more vulnerable. The Dusky Langur occurs in at least nine protected areas: Cantor’s Langur in Penang National Park in Malaysia; Reid’s Dusky Langur in Krau Wildlife Reserve and Taman Negara National Park in Malaysia; the Blond-tailed Langur in Khao Sok, Kaeng Krachan, Khao Sam Roi Yot, and Thale Ban national parks in Thailand; the Tarutao Langur in Taratau National Park in Thailand; and the Phangan Langur in Phangan National Park in Thailand. Population estimates are not available, but populations are believed to be stable or only slowly declining. Mainland subspecies are widespread and still common, while insular forms have restricted distributions. Survey work to determine the population and conservation status is needed, particularly for the insular forms.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB6FFBCFA3F6FE4FAFFF356.taxon	materials_examined	Burma, Arakan.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB6FFBCFA3F6FE4FAFFF356.taxon	discussion	1. playrei is a member of the obscurus species group, which also includes 7. obscurus, 1: crepusculus, and 1. barbei. Two subspecies recognized.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB6FFBCFA3F6FE4FAFFF356.taxon	distribution	Subspecies and Distribution. 1. p. phayrei Blyth, 1847 — E Bangladesh, NE India (Assam, Mizoram, and Tripura states), and W Myanmar (SE through Arakan to Pegu). I. p. shanicus Wroughton, 1917 — SW China (Yingjiang-Namting River and Tunchong-Homushu Pass districts in W Yunnan Province), and N & E Myanmar (Shan States and neighboring dry zone of N Myanmar).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB6FFBCFA3F6FE4FAFFF356.taxon	description	Descriptive notes. Head-body 42 - 60 cm (males) and 44 - 57 cm (females), tail 75 — 86 cm (males) and 72 - 80 cm (females); weight 5: 7.9 - 1 kg (males) and 6.4 - 7.5 kg (females). Phayre’s Langur is very similar to the Dusky Langur (7. obscurus) in overall appearance, butits legs, tail, and crown cap are not so contrastingly pale colored. Face is dark gray, except for white ocular rings and a depigmented area on mouth and nasal septum. The nominate subspecies (1. p. phayrei) is dark brown, buffy or gray-brown above, with sharply contrasting gray or whitish underparts. Its face is black, except for a fleshy-white muzzle and a white ring encircling each eye. There is no whorl behind its eyebrows. The “ Shan States Langur ” (1. p. shanicus) is generally fawn colored, with a trace of lightening on nape and hands. Its feet are slightly darker than the rest of its body. There is either a single tall whorl or a parting behind its brows, but no crest on its crown. The depigmented area around its mouth is very wide, and white aroundits eyes is restricted to innersides.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB6FFBCFA3F6FE4FAFFF356.taxon	biology_ecology	Habitat. Primary and secondary high-elevation evergreen, semi-evergreen, and mixed moist deciduous forests throughout most of its range. In some areas, it also occurs in sparse woodlands and bamboo forests. Phayre’s Langur prefers dense cover and bamboo thickets along hillsides and streams. It can also be found in tea plantations.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB6FFBCFA3F6FE4FAFFF356.taxon	food_feeding	Food and Feeding. Phayre’s Langur feeds mainly on leaves, flowers, seeds, sap, gum, figs, and bamboo shoots. In Bangladesh, mature leaves and young shoots form a major part of the diet (66 %), followed by flowers and buds (16 %) and fruits and seeds (14 %). Seasons influence dietary choice, with leaves dominating the diet in December (76 %) and the lowest preference for foliage (49 %) in April. Fruits and seeds are preferred mostly in June (57 %) when leaf consumption is low (15 %). Consumption of flower and buds peaks in April (41 %) and is lowest in January (9 %). Bamboo shoots form a significant proportion ofits diet.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB6FFBCFA3F6FE4FAFFF356.taxon	breeding	Breeding. Natal coat of Phayre’s Languris straw-colored. The gestation period of the nominate subspecies is 205 days, and the reproductive cycle is 28 days. Females are receptive throughout the ovarian cycle and during gestation. The average mating interval is 17 days. Female Phayre’s Langurs mate with subadult males during non-ovulatory periods and with alpha males during the periovulatory period. This reproductive strategy may confuse males regarding paternity and reduce the risk of infanticide from male aggression. Females mate in exchange for male security.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB6FFBCFA3F6FE4FAFFF356.taxon	activity	Activity patterns. Early mornings and late afternoons are peak foraging times for of Phayre’s Langurs in Bangladesh. They prefer to rest in certain trees such as the breadfruit tree (Artocarpus chaplasha, Moraceae) and Gmelina arborea (LLamiaceae) that provide good shade, and trees or bamboos such as Actinidia chinensis (Actinidiaceae), Melocanna baccifera, M. bambusoides and Bambusa tulda (Poaceae) or Acacia mangium (Fabaceae) that offer moderate shade.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB6FFBCFA3F6FE4FAFFF356.taxon	biology_ecology	Movements, Home range and Social organization. In Bangladesh, Phayre’s Langurs split into subgroups while foraging but communicate and maintain contact visually and with vocalizations. Group size has a significant effect on infant development. Infants develop more slowly in the larger groups. This phenomenon is attributed to intragroup competition for resources that exert a negative effect on individual energy and time budgets.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB6FFBCFA3F6FE4FAFFF356.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Endangered on The IUCN Red List, including both subspecies. Both subspecies have undergone declines of more than 50 % over the last three generations because of habitat loss and hunting. Phayre’s Langur is protected under Schedule I, Part I of the Indian Wildlife (Protection) Act, 1972; Schedule III of the Bangladesh Wild Life (Preservation) Order, 1973; and Category I of the Chinese Wildlife Protection Act. In Myanmar, it has been protected since 1994. Phayre’s Langur is commonly hunted for its gallstones, which are used in traditional medicine. In 2008, the total number of the Shan States Langur was estimated at fewer than 2500 individuals. Phayre’s Langur occurs in at least eleven protected areas: the nominate subspecies in Lawachara National Park and Rama-Kalenga Wildlife Sanctuary in Bangladesh and Dampa, Gumti, Sepahijala, and Trishna wildlife sanctuaries in India, and the Shan States Langur in Gaoligongshan, Daxueshan, and Tongbiguan national nature reserves in China and Pidaung Wildlife Sanctuary and Popa Mountain National Park in Myanmar. In India and Bangladesh, populations are small and severely fragmented and are affected by changes in land use such as conversion of forests to tea and timber plantations, logging for paper mills and charcoal production, livestock ranching, shifting agriculture and human settlement, and habitat degradation through harvesting of non-timber forest produce. In Bangladesh, Phayre’s Langur has declined by more than 80 % in the last 20 years because of tourism, plantations, exotic species replacing indigenous varieties, encroachments, and the construction of roads and railway tracks. In China, populations of Phayre’s Langur in Gaoligongshan Nature Reserve and Tongbiguan Nature Reserve are healthy. There is no information on its status in Myanmar.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB9FFBCFFE469E2F98EF556.taxon	distribution	Distribution. SW China (E of Salween River and S of Xishuanbanna, S Yunnan Province), S Myanmar, N Thailand (S to Raheng, and W to the coast of the Bay of Bengal), N & C Laos, and N Vietnam, the W limit is most likely the Salween River; S of the distribution of Phayre’s Langur (1. phayrer).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB9FFBCFFE469E2F98EF556.taxon	description	Descriptive notes. Head-body c. 51 cm (males) and c. 49 cm (females), tail ¢. 83 cm (males) and c. 82 cm (females); weight ¢. 6 - 9 kg (males) and c. 6 - 4 kg (females). Female Indochinese Gray Langurs are slightly smaller than males. Body and tail are light gray, and belly is slightly brighter and silvery gray. Forehead, cheeks, and hair around face is brown. Feet and hands are dark gray. Color of forearms merges into light gray color of body. Tail ends with a thin tassel. Head has no crest but long hairs form a whorl. Bare facial skin is dark gray, with full light eye rings and depigmented patches in the middle of upper and lower lips.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB9FFBCFFE469E2F98EF556.taxon	biology_ecology	Habitat. Primary and secondary evergreen and semi-evergreen forests, mixed moist deciduous forests, and also light woodland, bamboo-dominated areas, near plantations, and even in limestone regions. In Laos, it occurs in limestone (karst) forests. Depending on the region, the Indochinese Gray Langur is sympatric with members of the francoisi species group and the Red-shanked Douc (Pygathrix nemaeus).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB9FFBCFFE469E2F98EF556.taxon	food_feeding	Food and Feeding. The Indochinese Gray Langur is largely folivorous and also eats figs, bamboo shoots, seeds, flowers, and gums.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB9FFBCFFE469E2F98EF556.taxon	breeding	Breeding. Ovarian cycle of the Indochinese Gray Langur is ¢. 28 - 4 days, and females undergo an average of 3 - 6 cycles until conception. The average gestation period recorded for seven females was c. 205 days. Births occur throughout the year, but most are in November — April. Interbirth intervalsif the previous infant survives are c. 2 years.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB9FFBCFFE469E2F98EF556.taxon	activity	Activity patterns. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB9FFBCFFE469E2F98EF556.taxon	biology_ecology	Movements, Home range and Social organization. Groups of Indochinese Gray Langurs can be as large as 14 - 20 individuals, with 1 - 5 adult males and 2 - 12 adult females. Home ranges are 73 - 7 - 93 - 4 ha. Females disperse and regularly transfer among groups. After immigrating, they can quickly take over the alpha position in the female hierarchy. Population densities recorded for Nam Nao National Park and Phukhio Wildlife Sanctuary in Thailand were 23 - 38 ind / km?, or an average of 3 - 4 groups / km?.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB9FFBCFFE469E2F98EF556.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Endangered on The [UCN Red List (as 1. phayrei crepuscula). Although the Indochinese Gray Langur has a wide distribution, it is rarely seen, and populations can be expected to have declined severely in recent years. It is threatened by habitat loss and hunting throughout its range. It often develops large gallstones that are used in traditional Asian medicine. A global population estimate is not available. In China, Indochinese Gray Langurs have been extirpated from many areas and have fallen to less than 5400 individuals (this number includes also the “ Shan States Langur, ” 1. phayrei shanicus). Due to low densities and its presence in few isolated localities, the Indochinese Gray Langur is considered critically endangered in Vietnam. In Laos, it is rare with only a few records. The Indochinese Gray Langur occurs in Pu Mat, Ben En, Xuan Lien, and Pu Luong national parks and Muong Ne Nature Reserve in Vietnam; Phou Dendin, Nam Et-Phou Louey, Phou Khaokhoay, and Nam Kading national biodiversity conservation areas and Phou Gnouey Production Forest in Laos; Nam Nao National Park and Huai Kha Khaeng and Phu Khieo wildlife sanctuaries in Thailand; and Ailaoshan, Huanlianshan, Wuliangshan, Xishuangbanna, Fenshuiling, Niuluohe, Caiyanghe, and Lancangjiang national nature reserves in China.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB9FFBDFAEE6FE3F8C8FDA8.taxon	materials_examined	the type locality is supposed to be the village Cai Kien (21 ° 19 ° N, 107 ° 44 ’ E), or the Cai Kinh limestone massif 21 ° 45 ’ N, 107 ° 30 ’ E.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB9FFBDFAEE6FE3F8C8FDA8.taxon	discussion	These type localities are outside the known distribution of 1. poliocephalus, and itis proposed to restrict it to Cat Ba Island, Vietnam. In the past, 1. poliocephalus was mostly classified as a subspecies of 1. francoisi or placed as the nominate taxon together with 7. leucocephalus. It is a member of the francoisi species group, also known as the karst or limestone langurs. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB9FFBDFAEE6FE3F8C8FDA8.taxon	distribution	Distribution. Cat Ba I in Ha Long Bay, off N Vietnam.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB9FFBDFAEE6FE3F8C8FDA8.taxon	description	Descriptive notes. Head-body 49 - 59 cm (males) and 49 - 55 cm (females), tail 82 - 92 cm (males) and 82 - 89 cm (females); weight 8.9 - 5 kg (males) and 6.7 - 8 kg (females). Pelage of adult male and female Cat Ba Langursis a dark chocolate-brown, not black. Head and neck, down to shoulders, is bright golden-brown to yellowish-white, with head and crest being the lightest colored parts. Hairs on head form a crown crest; central hairs of crest are often brown tipped. A 5 cm-wide gray band runs from thighs to back, above root oftail. This band forms a V-shape, with the tip on lower back. Hairs of the band are dark brown, like the rest of back, but 5 — 10 mm of the tips are bright silver gray, giving the band a “ frosted ” appearance. Feet and hands have small yellow patches that are the same color as shoulders. Female Cat Ba Langurs have a skin patch in their pubic areas, with irregular pigmentation and pale white or yellowish hairs. They have very long hairs on their backs, with hair length up to 180 mm that form a back cape. All bare skin, including on face, ears, hands, and feet, is black. Cat Ba Langurs are born with a bright yellow-orange pelage. After c. 6 months, back and tail grow darker, and there is an increase in the contrast between head and shoulders and the back. A shine of silver-gray hair indicates beginning of the band running from thighs to back. Coat grows darker, and after 1 - 5 - 2 years, individuals have adult coloration.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB9FFBDFAEE6FE3F8C8FDA8.taxon	biology_ecology	Habitat. Forests on limestone (karst) hills. Cat Ba Langurs belong to the so-called “ limestone langurs. ” Especially in winter, groups use caves and crevices as sleeping places. The region is characterized as a humid, subtropical monsoon area, with pronounced wet and dry seasons. The rainy season lasts from May to September, with the heaviest rainfall occurring in July-August. Dry and cold seasons last from November to March. Mean annual rainfall is ¢. 1800 mm, and temperatures can drop to 0 ° C.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB9FFBDFAEE6FE3F8C8FDA8.taxon	food_feeding	Food and Feeding. Cat Ba Langurs are mainly folivorous, with leaves making up the highest percentage of their diets, and the remainder composed of shoots, fruits, flowers, and bark. Their preferred habitat is bush and shrub plant communities. Ninetyeight plant species from 42 families have been recorded in their diet.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB9FFBDFAEE6FE3F8C8FDA8.taxon	breeding	Breeding. Cat Ba Langurs are born throughout the year but with a peak in the beginning of the year. Females give birth to a single offspring.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB9FFBDFAEE6FE3F8C8FDA8.taxon	activity	Activity patterns. Cat Ba Langurs are diurnal, crepuscular, arboreal, and terrestrial. Sleeping places, mostly caves or crevices, are left before dawn. Peak feeding times are 07: 00 - 09: 00 h, 11: 00 - 12: 00 h, and 15: 30 - 17: 00 h. In the afternoon, groups move near cliffs with caves and remain in the vegetation. When they finally arrive at their sleeping cave, it is nearly dark (summer c. 18: 30 h and winter ¢. 17: 30 h). On average, 66 % of the day is spent resting, 15 % feeding, 11 % traveling, and 8 % engaging in social behavior (observations in April-May). Activity budgets of adults andjuveniles are similar, but infants spend 33 % oftheir daily activities engaged in social behavior, including playing, and they spend less time in resting.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB9FFBDFAEE6FE3F8C8FDA8.taxon	biology_ecology	Movements, Home range and Social organization. Groups of Cat Ba Langurs move systematically, not randomly, through their home range. Like other langurs of the “ limestone langur ” group, quadrupedalism is the most frequent form of locomotion on both rocks and in trees. Following a rest period, a group generally moves to a new location. During rest periods, group members usually stay close together. Some individuals are in close contact, while others are separated by 10 - 20 m. At feeding times, distances between individuals increase up to ¢. 100 m. The adult male in the group usually initiates movement. The male starts to move to a higher elevation and then sits down 100 - 300 m from other group members to observe the locality, sometimes for 15 - 30 minutes. If the group follows him, the male relocates to another observation point. They often use tops of cliffs to inspect the surrounding area, while the group below moves from one slope or valley to another. During walking, running, and leaping, Cat Ba Langurs display a concave downward arch of theirtails, as in the northern distributed species of the “ limestone langurs, ” Francois’s Langur (71. francoisi), the White-headed Langur (7. leucocephalus), and Delacour’s Langur (71. delacouri). This tail posture differs from the southerly distributed species, the Hatinh Langur (7 hatinhensis), the Laos Langur (7. laotum), and the Black Langur (7. ebenus). There is little information on home ranges of Cat Ba Langurs. Distances between observations of one group were only 1 - 2 km, and the home range was c. 30 ha. Groups are composed of an adult male, several adult females, and their offspring. Surplus males live alone or join together in bachelor groups. In the past, group size in an undisturbed population was 8 - 10 individuals. In 2000, an average group size of 5 - 8 individuals was recorded on Cat Ba Island. In 1992, the population density was estimated at 0 - 003 ind / km?.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB9FFBDFAEE6FE3F8C8FDA8.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Critically Endangered on The IUCN Red List. The Cat Ba Langur has been listed as one of the “ World's 25 Most Endangered Primates ” since 2000. It is protected by Vietnamese law. The current total population is only 65 - 70 individuals. Despite widely intact habitat, the population is fragmented into 5 - 7 groups that have very limited or no contact. The historical density of Cat Ba Langurs on the island is unknown. According to older villagers and early scientific visitors, they occurred in high numbers in the past. It is assumed that the entire island of Cat Ba (c. 140 km?) had previously been optimal habitat and could have supported at least 2400 - 2700 individuals. The population began to decrease in the 1960 s, and this decrease was further accelerated by an increase in human settlements in subsequent decades. Between 1970 and 1986, ¢. 500 - 800 Cat Ba Langurs were killed. Despite protection of most Cat Ba Langurs inside Cat Ba National Park, the population of 104 - 135 individuals in 2000 declined by roughly 50 % by 2005. Poaching for meat and traditional medicine has been the most severe threat. A preferred hunting practice has been to catch Cat Ba Langurs, mostly the whole group, inside a sleeping cave. Habitat disturbance and fragmentation have resulted from the growing human population on Cat Ba Island, in part because of the country’s designation of the island as a new tourism center for northern Vietnam. Rampantfires started by honey collectors are also threats. Tourism in the inadequately managed Cat Ba National Park in the past has directly affected the natural history of Cat Ba Langurs by direct disturbance to the habitat. In 2000, a captive breeding program was established at the Endangered Primate Rescue Center, Cuc Phuong National Park, Vietnam.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB8FFBEFA3C678CFB7AF4E6.taxon	materials_examined	Fusui County, Guangxi, China.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB8FFBEFA3C678CFB7AF4E6.taxon	discussion	T: leucocephalus is a member of the francois: species group, also known as the karst or limestone langurs. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB8FFBEFA3C678CFB7AF4E6.taxon	distribution	Distribution. S China, restricted to the counties of Fusui, Chongzuo, Ningming, and Longzhou in SW Guangxi Autonomous Region; it is bordered in the NW and N by the Zuojiang River, in the SW and S by the Mingjiang River, and to the SE by the Shiwan Mts.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB8FFBEFA3C678CFB7AF4E6.taxon	description	Descriptive notes. Head-body 55 - 62 cm (males) and 47 - 55 cm (females), tail 82 — 89 cm (males) and 77 - 82 cm (females); weight 8.9 - 5 kg (males) and 6.7 - 8 kg (females). Body coloration of the White-headed Langur is dark chocolate-brown. Head is white to ivory-colored, and its hairs form a crest, sometimes with brownish hairs in the tip. White color extends down to shoulders, upper back, and upper chest. White areas on hands, forearms, feet, and lower legs vary individually. Forearms in front are occasionally white, as are feet and lower legs. Tail is brownish at its base and from its tip is white halfto three-quarters of its length. In female White-headed Langurs, white sections on tail, arms, and legs diminish with age. This probably also occurs in males, but there are no long-term observations because of frequent replacement of males in a group. Differences in coloration have led to the assumption of hybridization with Francois’s Langur (7. francoisi), or that the taxon represents a partial albinistic form of Francois’s Langur. Female White-headed Langurs have a white pubic patch of depigmented skin, with white or yellowish-white hairs. All bare skin on face, ears, hands and feet, is black. White-headed Langurs are born with a bright yellow-orange pelage. Infants change to adult coloration during the first six months oflife.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB8FFBEFA3C678CFB7AF4E6.taxon	biology_ecology	Habitat. Limestone karst hills, covered with typical limestone forest. Most of the habitat of the White-headed Languris highly fragmented; limestone cliffs, 200 - 300 m high, are often surrounded by cultivated land with intensive agricultural activities. There are four different vegetation communities on these cliffs that are used differentially by White-headed Langurs. Caves and crevices are used as sleeping sites. The caves offer protection from predators and temperature extremes. Average rainfall in the area is c. 1000 mm / year. Average daily temperature is 22 - 1 ° C, with a maximum of 39 - 5 ° C and a minimum of — - 0 - 5 ° C.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB8FFBEFA3C678CFB7AF4E6.taxon	food_feeding	Food and Feeding. White-headed Langurs are mainly folivorous, with leaves making up 60 - 92 % of their diets; the rest being taken up by shoots, fruits, flowers, and bark. Diets changes during the year, but they always prefer young leaves. There are also differences in species composition of diets among groups and subpopulations. In Fusui Nature Reserve, White-headed Langurs feed on ¢. 50 of 164 available plant species, and leaves make up ¢. 90 % of the diet, of which 60 - 80 % is young leaves and the restis fruits, flowers, buds, and bark. In Bapen Nature Reserve, parts of at least 85 plant species are eaten, and leaves account for 91 - 4 % of the feeding records, of which 79 - 1 % are young leaves. One adult in captivity needed 590 - 750 g fresh leaves / day. White-headed Langurs also need water for digestion. In captivity, total daily water intake (water from food and drinking water) was 919 ml, or 0 - 15 ml / g body weight, with 0 - 12 ml / g from food and 0 - 03 ml / g from drinking. In the wild, White-headed Langurs lick leaves in the early morning, and drinking at stone hollows is assumed because they are the only places that might contain rainwater. Licking rock faces is a common activity, but it is uncertain if minerals, insects, or lichens are also taken.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB8FFBEFA3C678CFB7AF4E6.taxon	breeding	Breeding. Male White-headed Langurs reach maturity at c. 5 five years, females at 3 - 5 — 4 years. In Nonggang Nature Reserve, age offirst copulation for females was c. 49 months, and age of first birth was 57 - 68 months. The gestation period is c. 214 days. Births take place throughout the year, but they peak in the first months of the year. In Nonggang, 85 - 7 % of births are in December — March, with a peak in January of 28 - 6 %. Females give birth to a single offspring. Birth interval is 18 - 28 - 4 months, and lactation lasts 17 - 5 — 20 - 5 months. One female raises an average of 2 - 5 offspring during her lifetime. Infant mortality in the first year oflife is 156 %. Allomothering behavior mostly occurs during the first month of an infant’s life and gradually decreases from then on. In Fusui Nature Reserve, only groups in high-quality habitat have offspring. As habitat in the Reserve degraded rapidly, groups in low-quality habitat apparently stopped reproducing; this was attributed to the lower diversity and availability of food plants.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB8FFBEFA3C678CFB7AF4E6.taxon	activity	Activity patterns. The White-headed Languris diurnal, crepuscular, arboreal, and terrestrial. Daily activity starts with them leaving a sleeping cave at 05: 30 - 06: 10 h in summer and 06: 45 - 07: 00 h in winter. They enter their caves at 19: 45 - 20: 00 h in summer and 17: 30 - 17: 45 h in winter. Luminosity is 0 - 3 - 0 - 9 lux when they begin leaving their caves and 2: 6 - 7 - 5 lux when they enter them at night. A group may use at least three caves, each for a period of 2 - 4 days. During a year, White-headed Langurs spent an average of 11 - 5 hours / day outside and 12 - 5 hours / day inside their caves, accounting for 48 % and 52 % of their time, respectively. The peak in morning feeding occurs upon leaving caves. There are two feeding peaks per day, one in the morning (c. 05: 00 — 10: 00 h) and one in the afternoon (c. 16: 00 - 17: 00 h). Timing of feeding is considered to be the result of adaption to living in the karst area. On sunny days in summer, temperatures of the surfaces of bare rocks are very high, which results in a higher air temperature over karst areas. During this time, a long rest period at ¢. 10: 00 - 16: 00 h is common. In winter, White-headed Langurs keep the same regime and use the resting time to sunbathe on bare rocks. Different subpopulations obviously have different time budgets. In Bapen Nature Reserve, ¢. 50 % of the day is spent resting, 15 % each for feeding, traveling, and engaging in social behavior, and ¢. 5 % playing. In the Longling area, six groups spent an average of 70 % resting, 15 % feeding, and 10 % traveling. In summer, resting time is higher, up to ¢. 80 %, while feeding time is lower, ¢. 10 %. In winter, resting time is reduced to ¢. 50 %, and feeding time increases to 23 %. During a study in Hua Shan Nature Reserve in January-May, resting time was 33 - 6 %, feeding time 20 - 8 %, playing 20 - 2 %, moving 13 - 6 %, social behavior 7 - 3 %, vocalization 4 - 1 - %, and sexual activity 0 - 4 %.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB8FFBEFA3C678CFB7AF4E6.taxon	biology_ecology	Movements, Home range and Social organization. Quadrupedalism is the most frequently form of locomotion of White-headed Langurs on rocks and trees, and they spend ¢. 71 % of their time when traveling moving on rock surfaces. During walking, running, and leaping, a concave downward arch of the tail is exhibited, as in the northern species of the “ limestone langurs, ” Francois’s Langur and the Cat Ba Langur (1. poliocephalus). This posture differs from the southern species, the Hatinh Langur (1. hatinhensis), the Laos Langur (71. laotum), and the Black Langur (7. ebenus). Home ranges of White-headed Langurs are 16 - 47 ha (average 30 - 5 ha). Densities are 8 ind / km? in Fusui Nature Reserve and 17 ind / km? in Longgang. The typical social unit is a polygynous group, with one adult male, several adult females, and their immature offspring. Two adult males in a group are probably brothers or father and son. Females in polygynous groups remain in their natal home range, forming a new group following a group takeover by an invading male. Young males live together and form bachelor groups of up to four individuals. When males become sexually mature, they travel alone and try to take over a group or to split females from a group. Infanticide has been observed in several cases after takeover of a group by an invader. Average group size differs among subpopulations: 5 - 1 individuals in Fusui Nature Reserve, 9 - 1 in the Longgang area, 5 - 3 in Bapen Nature Reserve, and 10 - 9 in the well-protected Chongzuo Eco Park Nature Reserve. Group size is influenced by habitat quality.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFB8FFBEFA3C678CFB7AF4E6.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Critically Endangered on The IUCN Red List (as T. poliocephalus leucocephalus). The White-headed Langur is protected by Chinese law. The total area of its distribution comprises ¢. 3500 km? but the fragmented area ofits occurrence is only ¢. 200 km ®, and subpopulations occur in 16 patches. The largest patch in Ningming and Longzhou is 68 km? while in Chongzou and Fusui County, habitatis highly fragmented with six patches of ¢. 43 km * in Chongzuo and nine patches of ¢. 88 km? in Fusui. In 2002, the total remaining population of the White-headed Langur was only 580 - 620 individuals: ¢. 250 individuals in Fusui, 90 - 100 in Chongzuo, and 210 - 240 in Ningming and Longzhou. Some subpopulations decreased dramatically in the 1980 s and 1990 s. In Bapen Nature Reserve, the population decreased by ¢. 25 %. Poaching has been the primary cause of the most recent declines. Clearing land with fire, charcoal production, and burning off agricultural residue are activities that often lead to uncontrolled fires that escape into adjacent karst areas. Tree cutting for firewood has caused further degradation to the remaining habitat. Nevertheless, the population in Nonggang has been growing in the last few years. The population in Guangxi, south of the Zuojiang River, is separated from Francois’s Langur north of the river. Some Francois’s Langurs have been released in the area occupied by the White-headed Langur, resulting in some hybrids. First generation hybrids and their descendants have features more similar to Francois’s Langurs.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFBBFFBFFFE06E73FE27F9AE.taxon	materials_examined	Hoi Xuan, Vietnam.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFBBFFBFFFE06E73FE27F9AE.taxon	discussion	1. delacouri was formerly classified as a subspecies of 1. francoisi. It is a member of the Jfrancoisi species group, also known as the karst or limestone langurs. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFBBFFBFFFE06E73FE27F9AE.taxon	distribution	Distribution. NC Vietnam in Hoa Binh, Ha Nam, Ninh Binh and Thanh Hoa provinces.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFBBFFBFFFE06E73FE27F9AE.taxon	description	Descriptive notes. Head-body 52 - 62 cm (males) and 57 - 60 cm (females), tail 81 - 90 cm (males) and 84 - 89 cm (females); weight 7.5 - 8.5 kg (males) and 7.3 - 7.7 kg (females). Black and white body coloration of Delacour’s Langur is unique among the South-eastern Asian langurs. Except for the white pubic patch in females, both sexes have the same pelage color. Pubic patch is formed by an area of unpigmented skin in front of callosities with whitish hairs and has an individually different irregular shape. Black coloration of body is interrupted by a sharply demarcated line in the middle of the back thatis white and extending to a similar line above knees, giving it the appearance of wearing a pair of white “ shorts. ” Face is black, with elongated grayish-white cheek whiskers stretching from corners of mouth to ears. White hairs reach behind ears where they are numerous and dense and form a white patch. There is a forward-pointing crown crest, and ears stand out from head. The long bushy tail is different from all other Asian langurs. Hairs stick out at right angles from thetail, and fur oftail is bushy and reaches c. 10 cm in diameter close to the root so that tail appears carrot-like. Hips of female Delacour’s Langurs occasionally change color from white to yellow or light brown. White hairs on their pubic patch can also change color, which is a result of staining by sweat. Females with babies are especially susceptible to changing color in summer. Delacour’s Langurs are born with a bright yellow-orange pelage — brighter than in the closely related Francois’s Langur (7. francoisi), the Hatinh Langur (7. hatinhensis), and the Cat Ba Langur (71. poliocephalus). Contrasting with these species, bare facial skin of infant Delacour’s Langurs is more yellow. After c. 5 weeks from birth, infant hair on hands and feet changes from dark orange to black. After 4 - 6 months, pelage changes to brownish-black and dark gray shorts become visible, head changes to light brown, and about one-third of tail ends in a tassel. At 9 - 18 months, pelage changes to black and shorts are light gray. At 18 - 36 months, body and head change to black and shorts to white, and tail loses the tassel shape and changes to the carrot-like shape, with larger diameter to the root.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFBBFFBFFFE06E73FE27F9AE.taxon	biology_ecology	Habitat. Broadleaf evergreen forest in typical limestone karst. Delacour’s Langurs occur in areas with steep limestone outcrops. Their occurrence in this harsh substrate is probably for several reasons: to provide shelter for sleeping during cold winter nights and for water sources during a prolonged hot-dry period. In winter, the temperature can drop to 5 ° C and the temperature inside caves is higher. Water sources are essential. The bowl-like surface of the limestone can store water in small, shadowed, deep valleys for a long period, providing necessary water reserves. Hilly limestone areas without steep outcrops, even undisturbed by humans, are not populated by Delacour’s Langurs, probably because of the absence of surface water and such reservoirs. Delacour’s Langurs occur mostly in lower elevations up to ¢. 500 m, and only occasionally higher, up to 1000 m in Pu Luong Nature Reserve. There is no overlap in the distribution of Delacour’s Langurs with other species of the francoisi species group.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFBBFFBFFFE06E73FE27F9AE.taxon	food_feeding	Food and Feeding. Delacour’s Langur is among the most folivorous langurs. Leaves make up to 80 % of its diet: 60 % young leaves and only ¢. 20 % mature leaves. Unripe fruits (c. 10 %), shoots, flowers are eaten variably, depending on seasonal availabilities. Preferred leaves have a high protein-to-fiber ratio. Insects or other small animals are not eaten. About 30 % of the daily activity budget of Delacour’s Langur is used for feeding. The dietary diversity of Delacour’s Langurs is high relative to the number of plant and tree species used as food, but they do not focus on special limestone plant species. About 60 tree and plant species are used for food in the wild and 137 plant and tree species have been eaten in captivity. An adult individual Delacour’s Langur eats ¢. 1100 g / day. They are dependent on water for digestion. Daily water consumption depends on air temperature, amounting to 0 - 02 ml / g body weight at 25 ° C and 0 - 07 ml / g body weight by 32 ° C.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFBBFFBFFFE06E73FE27F9AE.taxon	breeding	Breeding. Male Delacour’s Langurs reach reproductive maturity at c. 5 years of age, females at four years. The gestation period is 180 - 196 days. Females give birth to a single offspring. The interbirth interval is 19 - 25 months. Infants in captivity are weaned at 19 - 24 months (likely less in the wild).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFBBFFBFFFE06E73FE27F9AE.taxon	activity	Activity patterns. Delacour’s Langurs are diurnal, crepuscular, arboreal, and terrestrial. Especially in disturbed areas, groups leave their sleeping sites before dawn and return at dusk or in complete darkness. About 60 % of the day is spent resting, 30 % feeding, 6 % socializing, and 4 % traveling.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFBBFFBFFFE06E73FE27F9AE.taxon	biology_ecology	Movements, Home range and Social organization. In limestone areas with prevailing steep cliffs, Delacour’s Langurs spend nearly 80 % of the time on rocks. Quadrupedalism is the most frequent form of locomotion on rocks and in trees, more than double that of climbing. Nearly double the amount of time climbing is spent on rocks than on trees. Terrestrialism does not adequately describe their locomotion; they are cliffclimbers. They leap only 6 % of the time, much less than other African and Asian colobines. There are differences in frequencies of climbing and leaping between substrates; they leap three times more frequently on trees than on rocks. When leaping between support types, they more frequently use trees as a landing substrate than rocks. During walking, running, and leaping, a concave downward arch of the tail is exhibited, as is found in the northern species of the “ limestone langurs, ” Francois’s, Cat Ba, and White-headed (7. leucocephalus) langurs. This tail posture differs from the southern species, the Hatinh, Laos (7. laotum), and Black (71. ebenus) langurs. There have been only a few observations regarding home range size because of the threedimensionality of the habitat. Home range sizes are ¢. 20 - 50 ha and depend on the sizes of areas with vegetation and bare rocks. The normal social unit is a polygynous group, with one adult male, several adult females, and their immature offspring. Two males in one group that appear to be adults are probably relatives, brothers or father and son. A group comprises, on average, ten individuals, one adult male, four adult females, two to three subadults and juveniles, and two to three infants. The largest group observed comprised 17 individuals, with one adult male, nine adult females, and their offspring. Subadult males leave their family group and travel alone or form bachelor groups with other males. Up to four individuals can form such a bachelor group. If these males grow to maturity, they try to split females from a group, or to take over a whole group. When replaced, an adult male travels mostly alone. Home ranges of the groups generally overlap, and there are occasional fights between males on the borders. Sometimes females transfer to another group. An area with a few very small groups (e. g. a male with only one or two females) probably testifies to hunting, which hinders the formation of normal social units.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFBBFFBFFFE06E73FE27F9AE.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Critically Endangered on The IUCN Red List. Delacour’s Langur has been listed as one of the “ World’s 25 Most Endangered Primates ” since 2000. It is protected by Vietnamese law. Its distribution is highly fragmented and restricted to limestone outcrops. The total distribution comprises ¢. 5000 km?, but their area of occurrence is limited to ¢. 400 km ®. The northern border of the distribution is formed by the Da River, and the southern border is most probably the Ma River. It is questionable whether Delacour’s Langur occurred south of the Ma River in the past. The population of Delacour’s Langur is less than 250 individuals, and it is fragmented into isolated subpopulations. During the 1990 s, 19 subpopulations were recorded. After 2000, at least nine were extirpated. Van Long Nature Reserve (Ninh Binh Province) harbors the largest population with more than 100 individuals, ¢. 50 % of the world’s total number. An estimated group of 30 - 40 exists in Pu Luong Nature Reserve (Thanh Hoa Province), and about the same number occurs in the unprotected Kim Bang area (Hoa Binh Province). Other subpopulations number less than 20 individuals. Poaching has been the biggest threat in the past. Delacour’s Langurs are prized for use in the preparation of a traditional medicine, “ monkey balm. ” Increasing limestone quarrying for cement production and building materials is a new threat and is shrinking habitat dramatically. A successful captive breeding program was established at the Endangered Primate Rescue Center, Cuc Phuong National Park, Vietnam, in 1996.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFBAFFBFFF3E638BF8C9F6E4.taxon	materials_examined	LLongzhou, China.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFBAFFBFFF3E638BF8C9F6E4.taxon	discussion	1. francoisi is a member of the francois species group, also known as the karst or limestone langurs. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFBAFFBFFF3E638BF8C9F6E4.taxon	distribution	Distribution. SC China (Chongqing Municipality and Guizhou and Guangxi provinces) and N Vietnam (Ha Giang, Cao Bang, Tuyen Quang, Bac Kan, and Thai Nguyen provinces). Francois’s Langur has the northernmost distribution of the langurs.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFBAFFBFFF3E638BF8C9F6E4.taxon	description	Descriptive notes. Head-body 51 - 63 cm (males) and 50 - 60 cm (females), tail 80 - 90 cm (males) and 74 - 90 cm (females); weight 6.4 - 7.9 kg (males) and 5.5 - 7.9 kg (females). Pelage of Francois’s Langur is glossy-black, with only a narrow tract ofslightly elongated white hairs running from corners of mouth along sides of face to upper edges of ears, in a sort of exaggerated handlebar moustache. They have a tall, pointed crest on their crowns. Face is jet black. There are no differences in coloration between males and females, except that females have a pubic patch with unpigmented skin and whitish or yellowish hairs in front of callosities. Pubic patch has an individually different irregular shape. Infants are born with a yellow-orange pelage but with individually variable blackish shades on back, hips, and tail. Facial skin is yellowish-brown. Color of hair starts to change to black at c. 3 weeks of age, and the infant is entirely black by six months of age.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFBAFFBFFF3E638BF8C9F6E4.taxon	biology_ecology	Habitat. Evergreen broadleaf, mostly limestone, forest. Steep limestone outcrops are the typical habitat, predominantly at elevations up to 600 m. Francois’s Langurs use caves and crevices on steep slopes and cliffs as shelters and sleeping sites. Minimum and maximum temperatures in the northern part of its distribution are 5 ° C and 40 ° C.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFBAFFBFFF3E638BF8C9F6E4.taxon	food_feeding	Food and Feeding. Francois’s Langurs are primarily folivorous, but fruits and seeds also make up a large part of their diet. Diets are variable among the populations. Ninety plant species provided food items for the population in Nonggang Nature Reserve, Guangxi, China. Ten species accounted for 60 % of the diet. In Fusui Nature Reserve, Guangxi, China, 37 plant species were recorded, and ten species accounted for 90 % of the diet. In Fusui, the diet was 94 % leaves and 6 % fruits, flowers, and stems, and in Nonggang, it was 53 % leaves (39 % young leaves, 14 % mature leaves), 17 % fruits, 14 % seeds, 8: 5 % flowers, and 7 % stems, roots, and bark. Francois’s Langurs lick rock surfaces and put pieces in their mouths, possibly for minerals.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFBAFFBFFF3E638BF8C9F6E4.taxon	breeding	Breeding. Male Francois’s Langurs reach sexual maturity at ¢. 5 years old, females at 3: 5 — 4 - 5 years. The menstrual cycle is 22 - 5 (+ 7 - 9) days. The gestation period is 184 (+ 15) days, and the interbirth interval is 19 - 1 (+ 2: 2) months. Females give birth to a single offspring.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFBAFFBFFF3E638BF8C9F6E4.taxon	activity	Activity patterns. Francois’s Langurs are diurnal, crepuscular, arboreal, and terrestrial. Caves and crevices are normal sleeping places. In the evening, they enter their sleeping cave, depending on the season, at 18: 30 - 20: 00 h and move out in the morning before dawn. There are two feeding peaks: 09: 00 - 11: 00 h and 16: 00 - 18: 00 h. Francois’s Langurs spend c. 43 - 52 % of the daytime resting, 23 - 25 % feeding, 14 - 17 % moving, 6 - 9 % playing, and 6 - 8 % grooming. Some activities change seasonally. In spring, when availabilities of young leaves are greater, they spend less time feeding and more time playing; in winter, more time is spent resting. In a small isolated population in Fusui Nature Reserve, Francois’s Langurs spend only 10 % of their day feeding in summer compared to 21 % in winter, because the abundance of leaves in the rainy season reduces feeding time. Juveniles play much more than adults: 15 % for juveniles compared to 3 % for adults.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFBAFFBFFF3E638BF8C9F6E4.taxon	biology_ecology	Movements, Home range and Social organization. In summer, Francois’s Langurs spend c. 60 % of their day in trees and 40 % on rocks; in winter, less time is spent on rocks. Leaping, quadrupedal walking, and climbing account for 92: 8 % of locomotory behavioral records, dropping or free-falling to lower levels of the canopy for 7 - 9 %, and brachiating for 2 - 1 %. During walking, running, and leaping, a concave downward arch of the tail is exhibited, as in other northern species of the “ limestone langurs, ” the Cat Ba Langur (71. poliocephalus), the White-headed Langur (7. leucocephalus), and Delacour’s Langur (71. delacouri). This posture differs from the southern species, the Hatinh Langur (7 hatinhensis), the Laos Langur (7. laotum), and the Black Langur (1 ebenus). There are no detailed studies on the home ranges of Francois’s Langurs, but some groups use less than 50 ha. The normal social unit is a unimale-multifemale group, with their offspring. The average group sizes were 5 - 7 - 7 - 5 individuals for several larger subpopulations with 18 - 75 total individuals in Guangxi. Average group size for all populations is 7 - 2 individuals.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFBAFFBFFF3E638BF8C9F6E4.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Endangered on The IUCN Red List. Francois’s Langur is protected by law in China and Vietnam. The population has decreased because of hunting for use in traditional medicine and is now highly fragmented. The total population declined in China during the last decade of the 20 " century by ¢. 70 % from an estimated 6000 - 7000 individuals to ¢. 1600. The population decline in Guangxi has been ¢. 85 %, to ¢. 300 individuals in 14 isolated populations. There are an estimated ten populations with 900 - 1000 individuals in Guizhou and four populations with ¢. 150 - 200 individuals in Chongqing. The largest population occurs in Mayanghe Nature Reserve, Guizhou, with ¢. 400 - 450 individuals. In Vietnam, Francois’s Langurs have been almost extirpated from their historical distribution; scattered populations exist in the northern provinces of Ha Giang, Cao Bang, Tuyen Quang, and Bac Kan. Populations in the provinces of Lang Son, Vinh Phuc, Lao Cai, and Yen Bai have already been extirpated. In Vietnam, there are only a few sightings of Francois’s Langurs that have been recorded in the last few decades. Based on interview surveys of local people, the total population in Vietnam is estimated to be less than 250 individuals, divided into at least ten subpopulations. Francois’s Langurs do not overlap with other “ limestone langurs. ” The population in Guangxi, China, is separated by the Zuojiang River from the White-headed Langur, which occurs to south. Some individual Francois’s Langurs have been released in the distribution of the White-headed Langur, which has resulted in some hybrids. First generation hybrids and their descendants have features most similar to Francois’s Langur.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFBAFFB8FA386C71FD34FA26.taxon	materials_examined	“ Indo China ” probably either Lai Chau or Fan Si Pan mountain chain (c. 22 ° 30 ’ N, 103 ° 50 ’ E), Vietnam.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFBAFFB8FA386C71FD34FA26.taxon	discussion	D. Brandon-Jones suspected the type locality of the holotype and sole recorded specimen was in the vicinity of Lai Chau, perhaps from the Fan Si Pan mountain chain, Vietnam. But there are strong indications that the specimen originated from central Laos. There is a contact zone, or even a small overlap, with 7. laotum in southern Phou Hi Poun National Protected Area, Laos. Black langurs in the southern part of Phou Hi Poun National Protected Area, Khammouane Province, and in limestone areas between this area eastwards on limestone blocks in the direction of Hin Namno, have a slightly different feature, with some white hairs around the pinna of the ears. The difference is hard to see in the field, but these animals also differ genetically from 1. ebenus and are possibly a distinct taxon. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFBAFFB8FA386C71FD34FA26.taxon	distribution	Distribution. EC Laos, in S Khammouane Province, close to the border to Vietnam and in scattered limestone blocks to the S into N Savannakhet Province to ¢. 16 ° 50 ° N, the speciesis also recorded in C Vietnam, Quang Binh Province (17 ° 59 ’ N, 105 ° 40 ’ E).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFBAFFB8FA386C71FD34FA26.taxon	description	Descriptive notes. Head — body 62 cm, tail 94 cm; weight 10 - 3 kg (measurements only exist from only a single male specimen). Pelage of adult Black Langurs is completely black, with a brownish tinge. Hairs on head form a crest. One male at the Endangered Primate Rescue Center, Vietnam, was completely black as a subadult, but as he aged to more than ten years, he developed a gray shine from corners of mouth to ears and down to his nape, in contrast to the Hatinh Langur (7. hatinhensis) that has a white line.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFBAFFB8FA386C71FD34FA26.taxon	biology_ecology	Habitat. Very steep and high limestone cliffs, most with sparse vegetation. Caves and crevices are used by Black Langurs as sleeping sites.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFBAFFB8FA386C71FD34FA26.taxon	food_feeding	Food and Feeding. There is no specific information for this species, but Black Langurs are known to be folivores.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFBAFFB8FA386C71FD34FA26.taxon	breeding	Breeding. The Black Langur has not been studied in the wild, but there are no differences between it and the Hatinh Langur in captivity. These langurs are born with a yellow-brown pelage but with individually variable parts of blackish shades on back, hips, and tail, which can be extensive in some individuals.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFBAFFB8FA386C71FD34FA26.taxon	activity	Activity patterns. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFBAFFB8FA386C71FD34FA26.taxon	biology_ecology	Movements, Home range and Social organization. Quadrupedalism is the most frequent form of locomotion. Due to their habitat, Black Langurs are much more terrestrial than Hatinh Langurs. The posture of the tail during running and jumping, a concave-upward tail inflection, corresponds with the other southern “ limestone langurs, ” the Hatinh Langur and the Laos Langur (7. laotum) and differs from the northern species, Francois’s Langur (71. francoist), Delacour’s Langur (1. delacouri), the White-headed Langur (7. leucocephalus), and the Cat Ba Langur (7. poliocephalus). The social structure of Black Langur groups is most probably very similar to that of the Laos Langur. Groups of 5 - 10 individuals have been seen in the wild.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFBAFFB8FA386C71FD34FA26.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Endangered on The IUCN Red List (under T. hatinhensis). Black Langurs are protected by law in Vietnam and Laos, but poaching in Laos is intensive. They occur in Hin Namno National Protected Area in Laos and Nui Giang Man Nature Reserve in Vietnam. Their inaccessible habitats provide for some protection.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFBDFFB8FFE36337FE2BF277.taxon	materials_examined	Ban Na Sao, SW French Laos, on the French side of the Mekong, in latitude about 17 ° 30 ° N.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFBDFFB8FFE36337FE2BF277.taxon	discussion	There is an overlap with 7. ebenus in Thakek and Gnomalath districts, south Phou Hi Poun National Protected Area. 1. laotum is a member of the francois: species group, also known as the karst or limestone langurs. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFBDFFB8FFE36337FE2BF277.taxon	distribution	Distribution. WC Laos, from S Bolikhamsal Province, probably with the Nam Kading River as the N barrier of the distribution, S to C Khammouane Province, at ¢. 17 ° 40 ’ N; there are also reports from the corridor to Nakai-Nam-Theun National Protected Area, and from Khamkeut District, close to Lak Sao (18 ° 13 ’ N, 104 ° 47 ’ E).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFBDFFB8FFE36337FE2BF277.taxon	description	Descriptive notes. Head — body 55 - 67 cm (males) and 46 - 54 cm (females), tail 85 — 90 cm (males) and 81 - 85 cm (females); weight 7 - 8 kg (males) and 6 - 7 kg (females). Body coloration of the Laos Langur is black, with a chocolate brown tinge, as in the Hatinh Langur (7. hatinhensis). Hairs on head, including cheeks and chin, are white except for crown, where black hairs form a crest. A stripe of black hairs runs from crest down to nape, sometimes connected with black pelage on back, sometimes interrupted with white hairs. Some black hairs form a scattered ring around face. Naked skin of face, ears, hands, and feet is jet-black. Females have a small irregular pubic patch, with depigmented skin and yellowish-white hairs.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFBDFFB8FFE36337FE2BF277.taxon	biology_ecology	Habitat. Steep limestone cliffs, with sparse vegetation. Patches of forest only remain on slopes and in valleys and canyons where the Laos Langur lives. About 50 % ofits distribution is rocky scrub, 30 % evergreen forest with shrubby vegetation, and ¢. 10 % mixed evergreen-deciduous forest.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFBDFFB8FFE36337FE2BF277.taxon	food_feeding	Food and Feeding. There is no specific information available for this species, but Laos Langurs are known to be folivores.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFBDFFB8FFE36337FE2BF277.taxon	breeding	Breeding. There is no information available for this species.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFBDFFB8FFE36337FE2BF277.taxon	activity	Activity patterns. Laos Langurs are diurnal, crepuscular, terrestrial, and arboreal. They are mainly terrestrial on limestone cliffs where theylive.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFBDFFB8FFE36337FE2BF277.taxon	biology_ecology	Movements, Home range and Social organization. There is no specific information available for this species, but Laos Langurs have been seen in groups of 5 - 10 individuals.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFBDFFB8FFE36337FE2BF277.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Vulnerable on The [UCN Red List. Poaching of Laos Langursstill occurs despite being protected by Laotian law. Habitat loss is currently no threat because the extremely steep and dry habitat where they live does not support agriculture. The Laos Langur occurs in Nam Kading and Phou Hi Poun national protected areas in Laos.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFBDFFB9FAEE6400F7E8FDAF.taxon	materials_examined	Xom Cuc (17 ° 56 ’ N, 105 ° 47 ° E) Ha Tinh Province (now in the Tuyen Hoa District, Quang Binh Province), Vietnam.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFBDFFB9FAEE6400F7E8FDAF.taxon	discussion	In the past, T. hatinhensis was placed as a subspecies of 1. francoisi or T. laotum. The taxonomic relationship to 7. ebenus and to another possibly distinct black langur in Laos is not fully clarified. 1. hatinhensis is a member of the francoisi species group, also known as the karst or limestone langurs. Monotypic.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFBDFFB9FAEE6400F7E8FDAF.taxon	distribution	Distribution. NC Vietnam (Quang Binh and Quang Tri province), and EC Laos (Khammouane Province). The records in other Vietnamese provinces (Nghe An, Thu Thien Hue, Gia Lai, Phu Yen) are errors.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFBDFFB9FAEE6400F7E8FDAF.taxon	description	Descriptive notes. Head-body 54 - 61 cm (males) and 55 - 58 cm (females), tail 83 — 95 cm (males) and 78 - 92 cm (females); weight 6 - 9 kg (males) and 6.2 - 8.2 kg (females). Male and female Hatinh Langurs are similar in coloration, except for the pubic patch of females. Pelage is black, with a brown tinge. A narrow tract of white hairs runs from mouth along cheeks to ears, extending and broadening behind ears down to nape. Color and extension of this white stripe is highly variable, from a broad and clear white band to a smaller more grayish line. Compared with Francois’s Langur (1. francoisi), whiskers of Hatinh Langurs do not have prolonged hairs on corners of mouth and a small white moustache connecting cheek stripes. Hairs on head form a crest that is bushier than in Francois’s Langur. As in other “ limestone langur ” species, female Hatinh Langurs have an irregular pubic patch, with depigmented skin and white or yellowish-white hair, extent of which is highly variable. Hatinh Langurs are born with a yellow-brown pelage but with individually variable patches of blackish shades on back, hips, and tail, which can be extensive in some individuals. Tail is already black at birth. Infants change to entirely black in less than six months. Facial skin is yellowish-brown and changes to black with pelage. Infant head color changes last. Occasionally, front part of head is yellowish-white until the age of two years, as is seen too in the Laos Langur (7. laotum).	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFBDFFB9FAEE6400F7E8FDAF.taxon	biology_ecology	Habitat. [Limestone karst with its typical vegetation. Hatinh Langurs prefer areas with dense forest cover. Dense forest is present in ¢. 90 % ofits known distribution. The core population occurs in one of the largest karst blocks in Indochina in Phong Nha-Ke Bang National Park in north-central Vietnam and Nakai-Nam Theun National Protected Area in east-central Laos. Caves and cliffs are used as sleeping sites, normally for many years if there is no disturbance or poaching. The heights of the caves or crevices on the cliffs vary from ¢. 20 - 60 m from their bases. Well-used sleeping sites are easily identified by the dark-yellow or brown stains on cliffs — marks of urine and feces.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFBDFFB9FAEE6400F7E8FDAF.taxon	food_feeding	Food and Feeding. Hatinh Langurs are highly folivorous. They eat leaves from 132 plant species, fruits from 26, roots from three, and bark from two. A group in a large semi-wild, 20 ha enclosure of primary forest in their natural environment in Phong Nha-Ke Bang National Park feeds mostly on leaves (81: 2 %) but also fruits (8: 7 %), tips of vines and climbers (8: 7 %), and flowers (1: 4 %). Daily food consumption of an adult averages 880 g. Daily water consumption of an adult during hot weather can be up to 500 ml / day.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFBDFFB9FAEE6400F7E8FDAF.taxon	breeding	Breeding. Male Hatinh Langurs reach maturity at c ¢. 5 years old and females at c. 4 years. The gestation period is 180 - 190 days. Females give birth to a single offspring. Births occur throughout the year but peak in January-March.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFBDFFB9FAEE6400F7E8FDAF.taxon	activity	Activity patterns. Hatinh Langurs are diurnal, crepuscular, and terrestrial, but mainly arboreal. There are no studies of their activity patterns in the wild. Hatinh Langurs in a captive population at the Endangered Primate Rescue Center, Vietnam, spend an average of 72 % of the day resting, 23 % feeding, 3 % engaged in social behavior, and 2 % moving. Their time budget is very similar to closely related species occurring in the wild, except that individuals travel more in the wild. In Phong Nha-Ke Bang National Park, Hatinh Langurs return to the vicinity of their sleeping sites, on average 65 minutes before sunset. They forage and play until they enter their sleeping caves, c. 14 minutes after sunset. They leave their caves and crevices before dawn at 05: 00 - 05: 30 h.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFBDFFB9FAEE6400F7E8FDAF.taxon	biology_ecology	Movements, Home range and Social organization. Quadrupedalism is the most frequent form of locomotion. The Hatinh Langur appears to be more arboreal than the other “ limestone langur ” species. Differing from Francois’s Langur, They hold their tails in a concave-upward inflection when walking and especially when running. In a large semi-wild, 20 ha area with natural vegetation, Hatinh Langurs spent 66 - 3 % of their time in trees, 17 - 7 % in scrub, 12: 4 % on fallen dead tree trunks, and 3: 6 % on rocks. There is no information about their home ranges in the wild. From 1995 to 1996, average group sizes of Hatinh Langurs in Phong Nha-Ke Bang National Park were 7 - 8 individuals. In 2007, average group size was 5 - 1 individuals (n = 23). Poaching may be the reason for the decrease. Groups normally have one male, three to four females, and their immature offspring. A group with 30 individuals has been seen.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
CE199B17FFBDFFB9FAEE6400F7E8FDAF.taxon	conservation	Status and Conservation. CITES Appendix II. Classified as Endangered on The IUCN Red List. The Hatinh Langur is protected by law in Vietnam and Laos. They occur in only two protected areas in Vietnam, Phong Nha-Ke Bang National Park in Quang Binh Province and Bac Huong Hoa Nature Reserve in Quang Tri Province. In Laos, they most probably occur only in Nakai-Nam Theun National Protected Area. Poaching is the main threat, mostly for traditional medicine. The main hunting method is to trap individuals inside their sleeping caves by covering the entrance with a net. Hunting with rifles has apparently diminished in Vietnam because of gun legislation controls and the confiscation of guns by local authorities. The largest population of Hatinh Langurs is most probably that in Phong Nha-Ke Bang National Park, with an estimated 1754 individuals. The population density is estimated at 0 - 5 groups / km? and 1 - 7 ind / km?. A captive breeding program was established at the Endangered Primate Rescue Center, Cuc Phuong National Park, Vietnam, and there have been successful births of Hatinh Langurs since 1996.	en	Russell A. Mittermeier, Anthony B. Rylands, Don E. Wilson (2013): Cercopithecidae. In: Handbook of the Mammals of the World – Volume 3 Primates. Barcelona: Lynx Edicions: 550-755, ISBN: 978-84-96553-89-7, DOI: http://doi.org/10.5281/zenodo.6867065
