Colobus guereza Ruppell 1835
Creators: name: Russell A. Mittermeier
name: Anthony B. Rylands
name: Don E. Wilson
Publisher: name: Lynx Edicions
Year: 2013
Subjects: Biodiversity
Taxonomy
Animalia
Chordata
Mammalia
Primates
Cercopithecidae
Colobus
Colobus guereza

description: 83.
Guereza
Colobus guereza
French: Colobe guéréza / German: Guereza / Spanish: Colobo guereza
Other common names: Eastern Black-and-white Colobus, Magistrate Colobus, Mantled Guereza; Djaffa Mountains Guereza/Neumann's Black-and-white Colobus (gallarum), Dodinga Hills Guereza (dodingae), Mau Forest Guereza (matschiel), Mount Kenya Guereza (kikuyuensis), Mount Kilimanjaro Guereza (caudatus), Mount Uaraguess Guereza/ Mount Uarges Guereza/Percival’s Black-and-white Colobus (percivali), Omo River Guereza (guereza), Western Guereza (occidentalis)
Taxonomy. Colobus guereza Ruppell, 1835,
Ethiopia, Gojjam and Kulla.
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.
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.
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.
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.
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.
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.
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.
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.
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 8year 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 1960s. 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.
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description: Published as part of Russell A. Mittermeier, Anthony B. Rylands & Don E. Wilson, 2013, Cercopithecidae, pp. 550-755 in Handbook of the Mammals of the World – Volume 3 Primates, Barcelona :Lynx Edicions on pages 699-701, DOI: 10.5281/zenodo.6867065