Mantled guereza (colobus)

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TAXONOMY

Suborder: Haplorrhini Infraorder: Simiiformes Superfamily: Cercopithecoidea Family: Cercopithecidae Subfamily: Colobinae Genus: Colobus Species: C. guereza Subspecies: C. g. caudatus, C. g. dodingae, C. g. kikuyuensis, C. g. matschiei, C. g. occidentalis, C. g. percivali Other names: C. guereza: C. abyssinicus, guereza, Abyssinan black-and white colobus, Abyssinian colobus, eastern black-and-white colobus, guereza colobus, guereza black-and-white colobus, magistrate colobus, magistrate black colobus, mantled guereza, white-mantled colobus; colobe à épaules blanches, colobe de l’Abyssinie, colobe guéréza (French); colobo rojo guereza (Spanish); abessinsk guerezavitsvansad guereza, östlig colobusapa, östlig svartvit guereza (Swedish); C. g. caudatus: Kilimanjaro guereza, Kilimanjaro colobus, Mt. Kilimanjaro guereza, white-tailed colobus, white-tailed guereza; C. g. dodingae: Dodinga Hills guereza; C. g. guereza: Abyssinian guereza, guereza, Omo River guereza, Rüppell’s guereza, typical guereza; C. g. kikuyuensis: Mt. Kenya guereza, Mt. Kenya colobus; C. g. matschiei: Matschie’s colobus, Mau Forest guereza; C. g. occidentalis: Congo guereza, magistrate colobus, western guereza; C. g. percivali: Mt. Uarges guereza, Mt. Uaraguess guereza, Percival’s black-and-white colobus. Groves (2005) lists 7 subspecies; C. g. caudatus, C. g. dodingae, C. g. guereza, C. g. kikuyuensis, C. g. matschiei, C. g. occidentalis, C. g. percivali, while formerly listing (Groves 2001) C. g. gallarum in addition to the aforementioned subspecies. Thus, C. g. gallarum (Djaffa Mountains guereza) persists in the literature on occasion but here is not considered a subspecies.

Conservation status: please search the IUCN Red List.

Life span: >30 years (captive) Total population: Unknown Regions: Equatorial Africa Gestation: 158 days Height: 61.5 cm (M), 57.6 cm (F) Weight: 9.3 to 13.5 kg (M), 7.8 to 9.2 kg (F)

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MORPHOLOGY

The guereza is a large black monkey with a white mantle, or ornamentation, and a tail tuft (Napier 1985). The body is mostly black, with the white mantle extending from the shoulder to the hip, connecting around the lower torso. The tail has a white tuft at its end which is variable in its extent along the length of the tail (Groves 2001). Subspecies are distinguished from one another by color variations in these features (Napier 1985). The face is surrounded by white hair, with bushy cheek hairs. There is a white stripe on the thigh (Groves 2001). In rare instances, almost entirely white individuals are reported from the west side of Mt. Kenya (Hull 1978).

Colobus guereza

C. g. guereza has a tail which is longer than the head and body combined, with the anterior half being gray and the tuft taking up about half of its length (DL Hull cited in Groves 2001; Groves 2001). The mantle is long and extends onto the back, becoming longer further back on the body (Groves 2001). C. g. dodingae has a tail which is substantially longer than the head plus the body and is white for only 40% of its distal end. This tail tuft is not particularly bushy. The hair is short and coarse and the mantle is only somewhat creamy in color, and does not expand up onto the dorsum (Groves 2001). C. g. matschiei has a tail which is significantly longer than the head and body with the tuft extending over less than half of its length. It has short hair with a yellowish mantle which does not extend onto the back but extends nearly to the white hairs around the face. The shoulders have some white (Groves 2001). The tail of C. g. occidentalis is longer than the head and body combined and the tuft extends only one-third of the tail from the distal end. The mantle is more a cream color than white and does not extend onto the back. The subspecies has some white on its shoulders (Groves 2001). C. g. percivali has a very long creamy yellow mantle and very long hair, extending longer than 40 cm (15.7 in) on the lower abdomen. The tail is as long as the head and body combined, with the white tuft extending over about two-thirds of its length (Groves 2001). C. g. kikuyuensis has a very large tail tuft, covering almost three quarters of its length and the anterior portions of the tail are grayish. The tail is roughly as long as the body and head together. The mantle is long, extends onto the back, and is over 40 cm (15.7 in) long on the lower abdomen. The thigh stripe is abbreviated (Groves 2001). C. g. caudatus has a longer mantle than C. g. kikuyuensis, its ventrum is less woolly, and over 80% of the tail is occupied by the tuft (DL Hull cited in Groves 2001; Groves 2001). Average weights for males fall between 9.3 and 13.5 kg (20.5-29.8 lbs) while for females, the range of averages is between 7.8 and 9.2 kg (17.2-20.3 lbs) (Napier 1985; WL Junger pers. comm. cited in Oates et al. 1994; Oates et al. 1994; Smith & Jungers 1997). Head and body length in males averages 61.5 cm (24.2 in) ranging from 54.3 to 69.9 cm (21.4-27.5 in). In females, it averages 57.6 cm (22.7 in) and ranges between 52.1 and 67.3 cm (20.5-26.5 in) (Napier 1985). Colobus guereza, like other Colobines, possesses a large and multi-chambered stomach which allows them to better digest plant fibers, including foliage. This ability to digest plant material is also assisted by bacteria in certain areas of the stomach. Together, these and other morphological adaptations allow the species to feed on large quantities of leaves (Oates & Davies 1994b). There is a range of dental sexual dimorphism throughout the subspecies of C. guereza, ranging from males having consistently larger teeth than females (e.g., C. g. caudatus), to similar dentition between the sexes (e.g., C. g. guereza), to instances where some female dentition is actually larger than male dentition (e.g, C. g. gallarum). In addition, in some cases, differences in canine size are reduced. These differences could be attributable to habitat and socio-sexual factors (Hayes et al. 1995). Guerezas primarily use quadrupedal locomotion and leaping to move through their environment, followed in frequency by climbing and other locomotor patterns (Mittermeier & Fleagle 1976; Morbeck 1977a; 1977b; Gebo & Chapman 1995; 2000). The species’ quadrupedal movement usually consists of bounds and gallops up and across large supports and when not moving, they will usually sit or recline (Mittermeier & Fleagle 1976; Gebo & Chapman 1995; 2000). Leaps are usually short and contribute to a generally horizontal or downward pattern of movement (Morbeck 1977a; Gebo & Chapman 1995). The species rarely is seen suspending and usually feeds above a support (Mittermeier & Fleagle 1976). While primarily arboreal, the species will descend to the ground to feed and to travel in cases were there are not suitable arboreal pathways (Oates 1977b). The guereza thumb is rudimentary and greatly reduced like most members of the colobus family (Napier 1985; Oates & Davies 1994b; Tague 2002). In captivity, guerezas have lived past thirty years (Weigl 2005).

RANGE

CURRENT RANGE MAPS (IUCN REDLIST): Colobus guereza Guerezas are distributed in a band across the center of Africa, from Nigeria and Cameroon east through the northern Democratic Republic of the Congo, through southern Sudan to Ethiopia, Kenya and Uganda and south into northern Tanzania (Napier 1985; Groves 2001). Within their range, guerezas are reported in the following countries; Cameroon, Central African Republic, Chad, Congo, Ethiopia, Gabon, Kenya, Nigeria, Rwanda, Sudan, Tanzania, Uganda and the Democratic Republic of the Congo (numerous sources compiled by Oates 1977b; Oates & Trocco 1983). C. g. guereza is found predominantly in Ethiopia, while C. g. occidentalis is found from Uganda west of the Nile, to southern Sudan, to Cameroon and south to Gabon. C. g. dodingae is only found in the southeastern Sudan. C. g. percivali is only found around Mt. Gargues in central Kenya. C. g. matschiei is found in western Kenya and south into northern Tanzania. C. g. kikuyuensis is found in central Kenya and C. g. caudatus is found in Tanzania, primarily in the region of Mount Kilimanjaro (Groves 2001).

HABITAT

Guerezas are tied to habitats that have trees and are present in both deciduous and evergreen forests (Oates 1977b; Oates et al. 1994; Lwanga 2006). They are found in forests and savanna woodlands within and to the north of the moist forests of central Africa, often extending into highland or montane forests (Oates et al. 1994). However, they are found in a variety of habitat types, including primary, secondary, riparian, gallery, and upland forest, especially those near rivers, lakes and with higher elevations (Dunbar & Dunbar 1974; Dunbar 1987; Oates 1977b). The species often prefers disturbed, secondary, or colonizing forests, and prefers degraded forests to old growth when both are available (Thomas 1991; Lwanga 2006). This preference is likely attributable to high species diversity of food trees in some secondary growth forests and may also be explainable in terms of milder chemical defenses in secondary growth species (Oates 1977a; Lwanga 2006). Other habitat types include moist lowland, medium-altitude and highland forests, rainforests, gallery forests, swamp forests and wooded grasslands (Oates 1977b; Dunbar 1987; Oates 1994; Fashing 2001b; Harris & Chapman 2007). Guereza will also occasionally visit swamps (Oates 1978). In addition, they can be found in high forests in mountainous areas, including altitudes up to 3300 m (10,826.8 ft) as well as areas under human use, such as eucalyptus plantations (likely visited to make up for nutritional deficiencies) (Dunbar & Dunbar 1974; Fashing et al. 2007; Harris & Chapman 2007).

Colobus guereza

Because of their wide distribution, both in location and altitude, average temperatures and rainfall figures can vary considerably between and even within study sites. At the Kibale National Park in western Uganda for example, daily annual temperatures in the moist evergreen forest range on average between 16.2°C and 23.3°C (61.2 °F and 73.9°F) with average rainfall values ranging between 157cm (61.8 in) and 175 cm (68.9 in) (Butynski 1990; Chapman et al. 2002). Being near the equator, only rainfall shows seasonal variation, peaking between March and May as well as between August or September and November (Oates 1977a; Chapman et al. 2002).

ECOLOGY

Leaves and fruit are the main foods of the guereza but the diet is quite variable as would be expected in a species with such a wide distribution and range of habitat types (Oates 1994; Fashing 2001b). While the species has historically been believed to be exclusively leaf-eaters, they are not obligate folivores (Oates 1994; Fashing 2001b). The 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 (Dunbar & Dunbar 1974; Oates 1994; Bocian 1997 cited in Kirkpatrick 1999; Fashing 1999; 2001; Harris & Chapman 2007). Fleshy fruits are usually consumed when unripe, with consumption being reduced as they fully ripen, likely to avoid competition with other primate species who prefer ripe fruit (Fashing 1999; Chapman et al. 2006; Harris & Chapman 2007). Often, while a number of species of plant are exploited, only several make up the majority of the diet at a specific site (Dunbar & Dunbar 1974; Clutton-Brock 1975; Oates 1977a; Dunbar 1987; Fashing 2001b; Preece 2006; Harris & Chapman 2007). The usual guereza pattern is to eat and select for young leaves, but in times of scarcity, to rely on mature leaves and fruit. However the use of mature leaves can vary widely across forests and between groups within the same forest (Oates 1977a; 1994; Preece 2006; Chapman et al. 2006; Harris & Chapman 2007). In addition, the possibility exists that fruit might be a preferred food item in some habitats (Fashing 2001b). Other foods consumed include bark and wood, seeds, flowers, petioles, lianas, arthropods, water-plants, concrete from buildings and soil (Oates 1978; Bocian 1997; Fashing 2001b; Chapman et al. 2006; Plumptre 2006; Fashing et al. 2007; Harris & Chapman 2007). Guerezas tend to eat foods with high protein-to-fiber ratios, and the availability of such foods is correlated with guereza biomass in a given habitat (Chapman et al. 2004). Finally, it is supposed that the species is flexible with its dietary needs, a factor which might allow its widespread distribution (Fashing 2001b). The diurnal guereza spends over half of its day resting (Oates 1977a; von Hippel 1996; Bocian 1997; Fashing 1999; 2001). The next most common activity is feeding, which also takes up a significant amount of time although sometimes locomotion is the second most common activity (Oates 1977a; von Hippel 1996; Bocian 1997; Fashing 1999; 2001). Other activities that occur far less frequently include vigilance, moving, grooming, greeting, clinging and play (Oates 1977a; von Hippel 1996; Fashing 1999). During the day, activities generally consist of periods of moving and feeding, punctuated by prolonged rest periods. This pattern is variable in number of times per day between study sites and specific days, but usually cycles around 3-5 times per day (Dunbar & Dunbar 1974; Oates 1977a; Bocian 1997). Guerezas leave the area around their sleeping trees one to several hours after sunrise and retire to sleeping trees by sunset (Bocian 1997). Groups occupy up to four nearby tall sleeping trees which are near food sources each night and try to avoid sleeping near other guereza groups (von Hippel 1998).

Colobus guereza

Home range is variable with study location, with full home range estimates ranging from just over .01 km² to 1 km² (.004 mi² to .4 mi²) with most estimates at the lower end of this range, usually under around .2 km² (.08 mi²) (Oates 1977a; 1977c; review of 11 studies recording home range in von Hippel 1996; Bocian 1997; Krüger et al. 1998; Fashing 1999; 2001a; Chapman & Pavelka 2005). Home ranges can overlap, often sharing home range area with several other groups (von Hippel 1996; Krüger et al. 1998; Fashing 2001a; Harris & Chapman 2007). In addition, there are core areas within the home range which are significantly smaller than the overall home range (Krüger et al. 1998; Fashing 2001a; Harris & Chapman 2007). When compared between study sites, groups are typically between 6 and 10 individuals and usually average under 12 individuals (Oates 1994; Krüger et al. 1998; Fashing 2007). In long-term studies, single-group day range averages were between 252 and 734 m (826.8 and 2408.1 ft), ranging as small as 62 m (203.4 ft) in a day to over 1360m (4461.9 ft) (Oates 1977a; Bocian 1997; Grimes 2000; Fashing 2001a). The main confirmed predator of the guereza is the crowned hawk-eagle (Stephanoaetus coronatus) (Oates 1977a; Struhsaker & Leakey 1990; Mitani et al. 2001; Fashing & Oates cited in Fashing 2007). A second predator of guereza is the chimpanzee (Pan troglodytes), which has been observed both successfully and unsuccessfully hunting the species (Suzuki 1975; Ihobe 2001). Leopards (Panthera pardus) are also potential predators of the species as guereza remains have been found in its scats. However, the species could also have fed upon carcasses deposited by eagles or resulting from falls (Hart et al. 1996). Other potential predators include other raptors such as Verreaux’s eagles (Aquila verreauxii) (Dunbar & Dunbar 1974). The guereza is often found living in sympatry with a number of other primate species. At the Ituri Forest, in the Democratic Republic of Congo, guerezas are part of one of the richest primate communities in the world. Here, they are found living in sympatry with Cercocebus albigena, C. galeritus, Cercopithecus hamlyni, C. l’hoesti, C. mitis, C. neglectus, C. pogonius, Colobus angolensis, Piliocolobus badius, Pan troglodytes, and Papio anubis (Thomas 1991). Other primates with which they are sympatric include but are not limited to Chlorocebus aethiops, Cercopithecus ascanius, Galago demidoff and Perodicticus potto (Bocian 1997; Grimes 2000; Stern & Goldstone 2005). In addition, infant guerezas will sometimes play with infant vervets (Chlorocebus aethiops) and the species will also sometimes associate with other sympatric species as well (Rose 1977; Chapman & Chapman 1996).

Content last modified: May 12, 2009

Written by Kurt Gron. Reviewed by Peter Fashing and Tara Harris.

Cite this page as: Gron KJ. 2009 May 12. Primate Factsheets: Guereza (Colobus guereza) Taxonomy, Morphology, & Ecology . <http://pin.primate.wisc.edu/factsheets/entry/guereza/taxon>. Accessed 2020 July 17.

SOCIAL ORGANIZATION AND BEHAVIOR

Guereza social groups are generally small and cohesive, generally ranging between 3 to 15 individuals with most groups at most study sites averaging within that range (Marler 1972; Oates 1977b; see von Hippel 1996 and Oates 1994 for reviews and Krüger et al. 1998 as an example). Groups of up to 23 have been studied and group sizes of up to 30-40 individuals are reported in the southern gallery forests of the Central African Republic, but even if the largest numbers are correct, groups this size are not typical of the species (Rose 1977; Fay 1985; von Hippel 1996; Fashing 2001a). Finally, group sizes tend to be larger in contiguous forest and smaller in riparian or smaller areas of forest (Oates 1977b; von Hippel 1996).

Colobus guereza

The usual composition of guereza groups includes one adult male (although several can be present as group size grows and in several populations, multimale groups are common), several adult females, and immature individuals (Oates 1977b; 1977c; Dunbar 1987; Oates 1994; von Hippel 1996; Oates et al. 2000; Fashing 2001c). The society appears to be matrilineally organized with females and immature individuals forming the closely bonded core of the group (Bocian 1997). Within multi-male groups, one male is dominant to the others and interactions between the adult males are aggressive, with some males eventually being forced out (Bocian 1997; Oates et al. 2000). When not a member of a group, males are seen either alone, or with other non-group males (pers. obs. cited in Fashing 2001c; Harris et al. 2009). Multi-male groups are unstable and often relapse into a single-male structure however it is unclear whether or not group size is limited by social constraints, environmental constraints, or some other factor (Dunbar & Dunbar 1976; Oates 1994; Chapman & Pavelka 2005). Group structure follows this general pattern throughout the guereza range (Oates 1994). In captivity, females do not exhibit a hierarchy of rank, but in the wild, some adult females have dominance relationships (Grunau & Kuester 2001; Harris 2005). Guerezas are not strictly territorial, although some populations participate in core area defense, especially during encounters with other groups that are usually aggressive in nature (von Hippel 1996; Harris 2006a; Fashing 2007). Adult males are significantly more active in inter-group encounters than females, although females do sometimes participate (von Hippel 1996; Fashing 2001c; Harris 2005; 2006b). Inter-group encounters function in mate defense and resource defense and often consist of chasing, increased activity, displays, vocalization but only rarely physical attacks (von Hippel 1996; Fashing 2001c; Harris 2005; 2006a). In addition, in at least one study, nearby guereza groups could be hierarchically ranked relative to one another, based on the outcome of encounters between themselves (Harris 2006b). While the group is resting, the male will often rest higher and away from the group, possibly to detect other groups and raptors (Oates 1977c). During group movements, the members of a guereza group stay near one another and follow a leading animal, often in single file and along a similar path, often changing leaders during each progression (Dunbar & Dunbar 1976; Oates 1977a; Fashing 1999) Within the guereza group, the majority of interactions are friendly and not agonistic, with allogrooming the main affinitive social behavior, especially between females (Oates 1977c; Bocian 1997; Grimes 2000; Harris 2005). Adult males rarely groom others within the group (Oates 1977c; unpub. data cited in Fashing 2007). When agonistic encounters within a group do occur, captive research shows that affiliative behavior occurs soon thereafter between former opponents, consisting of close proximity, grooming, embracing, mouth-opening, and hand touching (Björnsdotter et al. 2000). Captive data shows that facial expressions, vocalizations, and gestures/body positions in female guerezas do not signal dominance or subordination (Grunau & Kuester 2001). Female guerezas (at least in Kibale National Park, Uganda) do not typically disperse from their natal groups, although they may disperse when groups dissolve (Bocian 1997; Harris et al. 2009). One voluntary female dispersal event has also been recorded (Fashing 2007). In Kibale National Park, females may remain in their groups as adults even though their groups do not contain unrelated mates (due to long male tenure length) (Harris et al. 2009). Adult females within groups are often closely related but this is not true for all groups. Close adult female relatives may also occur among neighboring groups even though females are typically philopatric (Harris et al. 2009). Males typically disperse as subadults or adults. 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 (Marler 1972; Dunbar & Dunbar 1974; Oates 1974; Harris et al. 2009). Males that take over groups often have tenure lengths reaching five years or more (Harris et al. 2009). Some multi-male groups may consist of father-son pairs, but others apparently contain unrelated males (Harris et al. 2009). In Kibale 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 (Harris et al. 2009). It is uncertain as to whether or not females disperse from their natal groups as it has never been observed but they might transfer, particularly during disruptions in the composition of the group (Bocian 1997; pers. obs. cited in Fashing 2001c; Harris 2005). Dispersal is probably more common among males in guerezas than in females (unpub. data cited in Fashing 2007).

REPRODUCTION

Colobus guereza

Little is known about reproduction in the guereza (Harris & Monfort 2006). The species exhibits a harem-based, polygynous mating system and there is no strong seasonality of births (Oates 1974 cited in Oates 1994; Bocian 1997). About half of copulations are solicited by males and half by females (Harris & Monfort 2006). Solicitation behavior includes approaching the prospective partner and performing low-intensity mouth clicking or tongue-smacking (Hollihn 1973; Grimes 2000). To present, a female stands with her hindquarters facing a male with her arms bent, holding the body low without the tail raised. The copulation posture consists of the male grasping the female’s ankles and trunk from behind (Oates 1977c). Copulations usually occur within a group, but extra-group copulations have been observed in some populations (Fashing 2001c; Harris 2005). In addition, within multi-male groups, sometimes more than one male has mating access to females (von Hippel 1996). The ovarian cycle is around 24 days, with females receptive about 5 days before ovulation until 2-3 days after ovulation (Harris & Monfort 2006). Females exhibit no external signs of estrus (Oates 1994; Bocian 1997). Gestation length in wild individuals is around 158 days (5.2 mo) (estimated at 170 days (5.6 mo) in captivity) and the interbirth interval is around 22 months with a minimum of 16 months between births (Dunbar & Dunbar 1974; Rowell & Richards 1979; Harris & Monfort 2006). Birthrates are higher in habitats that are more forested (Dunbar 1987). Age at sexual maturity has been estimated at around 4 years old (48 months) in females and 6 years (72 months) in males (Oates 1977c).

PARENTAL CARE

At birth, guereza infants have white hair and pink to red skin in stark contrast with the black-and-white adults (Wooldridge 1971; Horwich & Manski 1975; Oates 1977c; Ackerman 1991). By three weeks old the face and ears start to darken and become grey (Oates 1977c). The natal coat and skin continue to darken and reach adult coloration around 3-4 months old with males attaining adult coloration faster than females (Oates 1977c; 1994; Ackerman 1991). Some grey coloration can persist above the ears on top of the head through 6-7 months after birth (Ackerman 1991). In captivity, six main behaviors are shown by females to infants, most frequently between birth and 4 weeks of age. These include the “kiss,” “two handed grasp,” “tail pull-up,” “urogenital licking,” “leg pull out,” and grooming (Horwich and Manski 1975). At birth in the wild, the infant is dependent on the mother, clinging to her during group movements. In the first 5 weeks of life in captivity, infant guerezas can’t control who is carrying them as they are not very mobile. After this age, they often choose their mothers and start actively returning to the mother at around 3 or 4 weeks old (Woolridge 1971; Horwich & Manski 1975). Infant guerezas will repetitively perform new motor skills once learned, often over and over for 15-20 minutes (Wolldridge 1971). Around 8-9 weeks old, the infant will start to eat vegetation, but in small quantities. Social play with other infant is also seen around this age. From the 14th to 17th week, the infant often plays, investigates, feeds, and moves on its own, but still clings to the mother during group movement. At 20 weeks, some movement independent of the mother occurs during group progressions. By 50 weeks old, the juvenile no longer clings to the mother and does not suckle anymore (Oates 1977c). In the wild and in captivity, infants are the focus of the attention of other members of the guereza group, especially females, and are often handled by these individuals, sometimes shortly after birth (Wooldridge 1971; Oates 1977c). This handling occurs for the first four months of the infant’s life but is curtailed thereafter, except for grooming. Such handling occurs most frequently in the first two weeks of the infant’s life and infants are often uncomfortable at being held by individuals other than their mothers (Oates 1977c). However, the mother allows others to take and handle the infant. In fact, in captive animals, more care may be given early in the infant’s life by non-mothers however this pattern does not necessarily follow in the wild (Horwich & Manski 1975; Oates 1977c). In captivity, males will play and groom only when with infants and their interest in the infant appears keyed toward their defense (Horwich & Manski 1975; Ackerman 1991). However, males are generally disinterested in infants at birth, only increasing their interest after the infant is 4-5 weeks old (Horwich & Manski 1975). Infants are always carried by the mother ventrally, with the infant grasping the mother’s fur (Woolridge 1971). In the wild, infant mortality in a small sample exceeded one third of infants dying before their pelage changed while after the color change, mortality was low (Dunbar & Dunbar 1974). High infant mortality is observed elsewhere in the wild as well (Oates 1977b). Infanticide has been observed in guerezas, usually committed by non-group or newly immigrated males (Onderdonk 2000; Harris & Monfort 2003).

COMMUNICATION

Guereza vocalizations and sounds can be roughly assigned to six basic categories “roaring,” “snorting,” “purring,” “cawing,” “squeaking and screaming,” and “tongue clicking” (Marler 1972; Oates 1977c). Perhaps the most frequent, characteristic, and studied vocalization of the guereza is high-intensity “roaring” bouts, which occur mainly during the night and at dawn, but can also occur during the day and at lower intensities when potential predators are nearby or during heightened arousal (Marler 1972; Oates 1974 cited in Oates 1977c; Oates 1977c; Bocian 1997; Fashing 1999; Oates et al. 2000). Roaring can be heard up to a mile (1.6 km) away and usually only one adult male per group will roar, even in multi-male groups (Marler 1972; Bocian 1997; Harris 2005). Roaring is contagious and functions in long-range communication; once an individual adult male starts roaring, other males in neighboring groups will also begin roaring, each calling from a different location (Marler 1972; Bocian 1997; Fashing 1999; Oates et al. 2000). Night roaring is spontaneous and most roaring is not associated with the nearby presence of other guereza groups (Oates 1977c). Variation in roaring among males can be explained by the ranking of their group relative to others and might also signal their fighting ability (Harris 2005; 2006b). Male roars have been shown to contain both honest and exaggerated information about body size (Harris et al. 2006).

Colobus guereza

“Snorting” is heard from all wild males in a group and sometimes from females in captivity. This type of vocalization can be uttered in response to potential predators, when approaching another animal before aggression, and is a potential alarm call (Marler 1972). “Purring” is heard prior to group movements and functions to help coordinate the group but also may be a low intensity alarm call (Marler 1972; Oates 1977c). “Cawing” is usually heard from females and infants, and occurs in situations of light distress. “Squeaks and screaming,” like “cawing” is heard from females and sub-adults. This vocalization is usually heard in times of strong distress, especially when infants feel threatened (Marler 1972). “Tongue-clicking,” while is used as an indicator of mild aggression, in both inter- and intra-group interactions. Its main function is likely as a threat (Marler 1972).

LISTEN TO VOCALIZATIONS

Colobus guereza: Male roar (African Primates at Home/Indiana University)

Seven facial expressions used in communication have been described in addition to ten body postures and movements, and six types of tactile communication (see Oates 1977c). Two of the most conspicuous displays performed by guereza adult males include flashy leaps during roaring, and “stiff legs,” often accompanying tongue clicking when two guereza groups meet (Oates 1977c).

Content last modified: May 12, 2009

Written by Kurt Gron. Reviewed by Peter Fashing and Tara Harris.

Cite this page as: Gron KJ. 2009 May 12. Primate Factsheets: Guereza (Colobus guereza) Behavior . <http://pin.primate.wisc.edu/factsheets/entry/guereza/behav>. Accessed 2020 July 17.

INTERNATIONAL STATUS

For individual primate species conservation status, please search the IUCN Red List. Also search the current scientific literature for primate conservation status (overall as well as for individual species), and visit CITES (Convention on International Trade in Endangered Species of Wild Fauna and Flora).

Conservation information last updated in 2009 follows, for comparison:

Colobus guereza

Guerezas are among the least threatened colobines, due to their propensity to travel on the ground as well as to use dry and gallery forest (Oates & Davies 1994a). Guerezas are one of the few primates that are generally considered able to cope with habitat degradation, are one of the primates which is least affected by it, and generally do well in degraded forests (Chapman et al. 2000; von Hippel et al. 2000; Fashing 2002). However, there are conflicting views on the effects of degradation on the species, even within the same site, with some researchers viewing population declines, while others noting increases (see von Hippel et al. 2000; Fashing 2002).

CONSERVATION THREATS

Threat: Human-Induced Habitat Loss and Degradation

Forest over-use and clearing for many uses leads to a decline in guereza numbers. Clearing of forest for firewood by locals contributes to population declines but commercial logging poses a larger threat. Such commercial uses include wood clearing for brewing beer, fuel, gin distillation, and for charcoal production (Chapman et al. 2007). Also, even if not cleared, there are a number of ways guereza habitat can be degraded. These include degradation due to, the cutting of trees for timber, charcoal, firewood, tools and other uses, the extraction of gold, livestock traffic, road building, agriculture, vandalization, poaching, extraction of plant foods, and the stripping of bark off of trees for medicine (Oates 1977b; von Hippel et al. 2000; Fashing 2004). However, guerezas sometimes respond positively to logging, and are sometimes more abundant and are found in higher densities in logged areas than unlogged, possibly due to an increase in preferred food trees after logging (Oates 1977b; Skorupa 1986; Oates & Davies 1994a; Plumptre & Reynolds 1994; Chapman et al. 2000). Complete forest clearance undoubtedly affects the species in a negative way and causes decreases in overall population. This was the case in western Uganda where the clearing of forest fragments caused a reduction in guereza numbers by over 50% over an eight year period (Chapman et al. 2007). In addition, sometimes forest is cleared for agriculture and replanted with coniferous trees which are unsuitable for guerezas.

Threat: Invasive Alien Species

Human activities can increase the concentration of elephants in guereza habitats, which occurred at Kabalega National Park, Uganda. In turn, high concentrations of elephants destroy guereza habitat and threaten populations (Oates 1977b).

Threat: Harvesting (hunting/gathering)

Hunting can also be a threat to guereza populations, but it is variable in its occurrence throughout the guereza range. Firearms as well as traditional weapons are used to hunt the species (Oates 1977b). In Gabon for example, hunted populations showed an 88% reduction in numbers versus unhunted populations (Lahm 2001). Elsewhere in its range, hunting is also common, as is the case in the northern areas of the Republic of Congo, where guerezas sold as bushmeat fetch between 4-9 USD (Eves & Ruggiero 2000). However, in Uganda, primates are not usually hunted for meat (Plumptre 2006). Historically, a large commercial trade in guereza skins has existed, especially during the 19th century but continued into recent times, often for fashion or the tourist trade (see Oates 1977 for a history). Not surprisingly, this represents a significant threat to the species.

Threat: Persecution

Crop-raiding by guerezas does occur, albeit at low levels (Naughton-Treves 1996; 1998). Thus, the potential for persecution as a result of conflict with humans does exist. Oates (1977b) comments that the killing of guerezas as crop-raiders could provide skins for commercial sale.

Threat: Changes in Native Species Dynamics

The African cherry tree (Prunus africana), a sometimes favored food for guerezas, has exhibited a notable decline across sub-Saharan Africa. While predominantly due to the harvesting of its bark for medicines, at least some of its deaths could be due to other factors, such as disease, insects, nutrient deficiency, or climate (Fashing 2004). The decline of this species negatively affects the guereza populations that rely upon it.

LINKS TO MORE ABOUT CONSERVATION

CONSERVATION INFORMATION

No current links for Colobus guereza
Links for all species

CONSERVATION NEWS

Bush-meat fuelling secretive trade in Guinea-Bissau’s primates (Ecologist; September 29, 2010)
Forest loss occurring around Kibale National Park in Uganda (Mongabay; June 28, 2010)
Logging Changed Ecological Balance For Monkeys, Damaged Health (ScienceDaily; June 15, 2005)
Links for all species

ORGANIZATIONS INVOLVED IN Colobus guereza CONSERVATION

Content last modified: May 12, 2009

Written by Kurt Gron. Reviewed by Peter Fashing and Tara Harris.

Cite this page as: Gron KJ. 2009 May 12. Primate Factsheets: Guereza (Colobus guereza) Conservation . <http://pin.primate.wisc.edu/factsheets/entry/guereza/cons>. Accessed 2020 July 17.

The following references were used in the writing of this factsheet. To find current references for Colobus guereza, search PrimateLit.

REFERENCES

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Colobus guereza

Chapman CA, Pavelka MSM. 2005. Group size in folivorous primates: ecological constraints and the possible influence of social factors. Primates 46(1):1-9.

Chapman CA, Balcomb SR, Gillespie TR, Skorupa JP, Struhsaker TT. 2000. Long-term effects of logging on African primate communities: a 28-year comparison from Kibale National Park, Uganda. Conserv Biol 14(1):207-17. Chapman CA, Chapman LJ. 1996. Mixed-species primate groups in the Kibale Forest: ecological constraints on association. Intl J Primatol 17(1):31-50. Chapman CA, Naughton-Treves L, Lawes MJ, Wasserman MD, Gillespie TR. 2007. Population declines of colobus in western Uganda and conservation value of forest fragments. Intl J Primatol 28(3):513-28. Chapman CA, Chapman LJ, Naughton-Treves L, Lawes MJ, McDowell LR. 2004. Predicting folivorous primate abundance: validation of a nutritional model. Am J Primatol 62(2):55-69. Chapman CA, Chapman LJ, Cords M, Gathua JM, Gautier-Hion A, Lambert JE, Rode K, Tutin CEG, White LJT. 2002. Variation in the diets of Cercopithecus species: differences within forests, among forests, and across species. In: Glenn ME, Cords M, editors. The guenons: diversity and adaptation in African monkeys. New York:Kluwer Acad/Plenum Pub. p 325-50. Clutton-Brock TH. 1975. Feeding behaviour of red colobus and black and white colobus in east Africa. Folia Primatol 23:165-207. Dunbar RIM, Dunbar EP. 1976. Contrasts in social structure among black-and-white colobus monkey groups. Anim Behav 24:84-92. Dunbar RIM, Dunbar EP. 1974. Ecology and population dynamics of Colobus guereza in Ethiopia. Folia Primatol 21:188-208. Dunbar RIM. 1987. Habitat quality, population dynamics, and group composition in colobus monkeys (Colobus guereza). Intl J Primatol 8(4):299-329. Eves HE, Ruggiero RG. 2000. Socioeconomics and the sustainability of hunting in the forests of northern Congo (Brazzaville). In: Robinson JG, Bennett EL, editors. Hunting for sustainability in tropical forests. New York: Colombia U Pr. p 427-54. Fashing PJ. 2001a. Activity and ranging patterns of guerezas in the Kakamega Forest: intergroup variation and implications for intragroup feeding competition. Intl J Primatol 22(4):549-77. Fashing PJ. 2007. African colobine monkeys: patterns of between-group interaction. In: Campbell CJ, Fuentes A, MacKinnon KC, Panger M, Bearder SK, editors. Primates in perspective. New York: Oxford U Pr. p 201-24. Fashing PJ. 1999b. The behavioral ecology of an African colobine monkey: diet, range use, and patterns of intergroup aggression in eastern black and white colobus monkeys (Colobus guereza). PhD dissertation, Columbia University. Fashing PJ. 2001b. Feeding ecology of the guerezas in the Kakamega Forest, Kenya: the importance of Moraceae fruit in their diet. Intl J Primatol 22(4):579-609. Fashing PJ, Dierenfeld ES, Mowry CB. 2007. Influence of plant of soil chemistry on food selection, ranging patterns, and biomass of Colobus guereza in Kakamega Forest, Kenya. Int J Primatol 28(3):673-703. Fashing PJ. 2001c. Male and female strategies during intergroup encounters in guerezas (Colobus guereza): evidence for resource defense mediated through males and a comparison with other primates. Behav Ecol Sociobiol 50(3):219-30. Fashing PJ. 2004. Mortality trends in the African cherry (Prunus africana) and the implications for colobus monkeys (Colobus guereza) in Kakamega Forest, Kenya. Biol Conserv 120(4):449-59. Fashing PJ. 2002. Population status of black and white colobus monkeys (Colobus guereza) in Kakamega Forest, Kenya: are they really on the decline? Afr Zool 37(2):119-26. Fay JM, 1985. Range extensions for four Cercopithecus species in the Central African Republic. Prim Conserv 6:63-8. Gebo DL, Chapman CA. 2000. Locomotor behavior in Ugandan monkeys. In: Whitehead PF, Jolly CJ, editors. Old world monkeys. Cambridge (UK): Cambridge U Pr. p 480-95. Gebo DL, Chapman CA. 1995. Positional behavior in five sympatric old world monkeys. Am J Phys Anth 97(1):49-76. Grimes KH. 2000. Guereza dietary and behavioural patterns at the Entebbe Botanical Gardens. MA thesis, University of Calgary. 146 p. Groves C. 2005. Order primates. In: Wilson DE, Reeder DM, editors. Mammal species of the world: a taxonomic and geographic reference, third edition, volume 1. Baltimore (MD): Johns Hopkins U Pr. p 111-84. Groves C. 2001. Primate taxonomy. Washington DC: Smithsonian Inst Pr. 350 p. Grunau T, Kuester J. 2001. Dominance style in female guerezas (Colobus guereza Rüppell 1835). Primates 42(4):301-7. Harris TR, Monfort SL. 2003. Behavioral and endocrine dynamics associated with infanticide in a black and white colobus monkey (Colobus guereza). Am J Primatol 61(3):135-42. Harris TR. 2006a. Between-group contest competition for food in a highly folivorous population of black and white colobus monkeys (Colobus guereza). Behav Ecol Sociobiol 61(2):317-29. Harris TR, Fitch WT, Goldstein LM, Fashing PJ. 2006. Black and white colobus monkey (Colobus guereza) roars as a source of both honest and exaggerated information about body mass. Ethology 112(9):911-20. Harris TR, Monfort SL. 2006. Mating behavior and endocrine profiles of wild black and white colobus monkeys (Colobus guereza): toward an understanding of their life history and mating system. Am J Primatol 68(4):383-96. Harris TR, Caillaud D, Chapman CA, Vigilant L. 2009. Neither genetic nor observational data alone are sufficient for understanding sex-biased dispersal in a social-group-living species. Mol Ecol 18:1777-90. Harris TR. 2006b. Within- and among-male variation in roaring by black and white colobus monkeys (Colobus guereza): What does it reveal about function? Behaviour 143(2):197-218. Harris TR. 2005. Roaring, intergroup aggression, and feeding competition in black and white colobus monkeys (Colobus guereza) at Kanyawara, Kibale National Park, Uganda. PhD dissertation, Yale University. 253 p. Harris TR, Chapman CA. 2007. Variation in diet and ranging of black and white colobus monkeys in Kibale National Park, Uganda. Primates 48(3):208-21. Hart JA, Katembo M, Punga K. 1996. Diet, prey selection and ecological relations of leopard and golden cat in the Ituri Forest, Zaire. Afr J Ecol 34:364-379. Hayes VJ, Freedman L, Oxnard CE. 1995. The differential expression of dental sexual dimorphism in subspecies of Colobus guereza. Intl J Primatol 16(6):971-96. von Hippel FA. 1996. Interactions between overlapping multimale groups of black and white colobus monkeys (Colobus guereza) in the Kakamega Forest, Kenya. Am J Primatol 38(3):193-209. von Hippel FA, Frederick H, Cleland E. 2000. Population decline of the black and white colobus monkey (Colobus guereza) in the Kakamega Forest, Kenya. Afr Zool 35(1):69-75. von Hippel FA. 1998. Use of sleeping trees by black and white colobus monkeys (Colobus guereza) in the Kakamega Forest, Kenya. Am J Primatol 45(3):281-90. Hollihn U. 1973. Remarks on the breeding and maintenance of colobus monkeys Colobus guereza, proboscis monkeys Nasalis larvatus and douc langurs Pygathrix nemaeus in zoos. Intl Zoo Ybk 13:185-8. Horwich RH, Manski D. 1975. Maternal care and infant transfer in two species of Colobus monkeys. Primates 16(1):49-73. Hull DB. 1978. Aberrations in the coat colour patterns of black and white Colobus monkeys. E Afr Wildl J 16:21-7. Ihobe H. 2001. Hunting attempt by chimpanzees on Abyssinian colobus at the Kalinzu Forest, Uganda. Pan Afr News 8(2):31-2. Kirkpatrick RC. 1999. Colobine diet and social organization In: Dolhinow P, Fuentes A, editors. The nonhuman primates. Mountain View (CA): Mayfield Pub Co. p 93-105. Krüger O, Affeldt E, Brackmann M, Milhahn K. 1998. Group size and composition of Colobus guereza in Kyambura Gorge, southwest Uganda, in relation to chimpanzee activity. Intl J Primatol 19(2):287-97. Lahm SA. 2001. Hunting and wildlife in northeastern Gabon: why conservation should extend beyond protected areas. In: Weber W, White LJT, Vedder A, Naughton-Treves L, editors. African rain forest ecology and conservation: an interdisciplinary perspective. New Haven (CT): Yale U Pr. p 344-54. Lwanga JS. 2006. Spatial distribution of primates in a mosaic of colonizing and old growth forest at Ngogo, Kibale National Park, Uganda. Primates 47(3):230-8. Marler P. 1972. Vocalizations of east African monkeys II: black and white colobus. Behaviour 42:175-97. Mitani JC, Sanders WJ, Lwanga JS, Windfelder TL. 2001. Predatory behavior of crowned hawk-eagles (Stephanoaetus coronatus) in Kibale National Park, Uganda. Behav Ecol Sociobiol 49(2-3):187-95. Mittermeier RA, Fleagle JG. 1976. The locomotor and postural repertoires of Ateles geoffroyi and Colobus guereza, and a reevaluation of the locomotor category semibrachiation. Am J Phys Anth 45:235-56. Morbeck ME. 1977a. Leaping, bounding and bipedalism in Colobus guereza: a spectrum of positional behavior. Ybk Phys Anth 20:408-20. Morbeck ME. 1977b. Positional behavior, selective use of habitat substrate and associated non-positional behavior in free-ranging Colobus guereza (Rüppel, 1835). Primates 18(1):35-58. Napier PH. 1985. Catalogue of primates in the British museum (natural history) and elsewhere in the British Isles, part III: family Cercopithecidae, subfamily Colobinae. London: British Museum (Natural History). 111 p. Naughton-Treves L. 1998. Predicting patterns of crop damage by wildlife around Kibale National Park, Uganda. Conserv Biol 12(1):156-68. Naughton-Treves L. 1996. Uneasy neighbors: wildlife and farmers around Kibale National Park, Uganda. PhD dissertation, University of Florida. 175 p. Oates JF, Davies AG. 1994a. Conclusions: the past, present and future of the colobines. In: Davies AG, Oates JF, editors. Colobine monkeys: their ecology, behaviour and evolution. Cambridge (UK): Cambridge U Pr. p 347-58. Oates JF, Davies AG, Delson E. 1994. The diversity of living colobines. In: Davies AG, Oates JF, editors. Colobine monkeys: their ecology, behaviour and evolution. Cambridge (UK): Cambridge U Pr. p 45-73. Oates JF. 1974. The ecology and behaviour of the black-and-white colobus monkey (Colobus guereza Rüppell) in East Africa. PhD dissertation, University of London. Oates JF, Bocian CM, Terranova CJ. 2000. The loud calls of black-and-white colobus monkeys: their adaptive and taxonomic significance in light of new data. In: Whitehead PF, Jolly CJ, editors. Old world monkeys. Cambridge (UK): Cambridge U Pr. p 431-52. Oates JF. 1994. The natural history of African colobines. In: Davies AG, Oates JF, editors. Colobine monkeys: their ecology, behaviour and evolution. Cambridge (UK): Cambridge U Pr. p 75-128. Oates JF. 1977a. The guereza and its food. In: Clutton-Brock TH, editor. Primate ecology: studies of feeding and ranging behaviour in lemurs, monkeys and apes. London: Acad Pr. p 275-321. Oates JF. 1977b. The guereza and man. In: Rainier III (Grimaldi) Prince of Monaco, Bourne GH, editors. Primate conservation. New York: Academic Pr. p 419-67 Oates JF. 1977c. The social life of a black-and-white colobus monkey, Colobus guereza. Z Tierpsychol 45:1-60. Oates JF, Trocco TF. 1983. Taxonomy and phylogeny of black-and-white colobus monkeys. Folia Primatol 40(1-2):83-113. Oates JF. 1978. Water-plant and soil consumption by guereza monkeys (Colobus guereza): a relationship with minerals and toxins in the diet? Biotropica 10(4):241-53. Oates JF, Davies AG. 1994b. What are the colobines? In: Davies AG, Oates JF, editors. Colobine monkeys: their ecology, behaviour and evolution. Cambridge (UK): Cambridge U Pr. p 1-9. Onderdonk DA. 2000. Infanticide of a newborn black-and-white colobus monkey (Colobus guereza) in Kibale National Park, Uganda. Primates 41(2):209-12. Plumptre AJ. 2006. The diets, preferences, and overlap of the primate community in the Budongo Forest reserve, Uganda: effects of logging on primate diets. In: Newton-Fisher NE, Notman H, Paterson JD, Reynolds V, editors. Primates of western Uganda. New York: Springer. p 345-71. Plumptre AJ, Reynolds V. 1994. The effect of selective logging on the primate populations in the Budongo Forest Reserve, Uganda. J Appl Ecol 31(4):631-41. Preece GA. 2006. Factors influencing variation in the population densities of Colobus guereza within selectively logged forest at the Budongo Forest Reserve: the importance of lianas during a subsistence diet. In: Newton-Fisher NE, Notman H, Paterson JD, Reynolds V, editors. Primates of western Uganda. New York: Springer. p 23-43 Rose MD. 1977. Interspecific play between free ranging guerezas (Colobus guereza) and vervet monkeys (Cercopithecus aethiops). Primates 18(4):957-64. Rowell TE, Richards SM. 1979. Reproductive strategies of some African monkeys. 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Content last modified: May 12, 2009

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	Colobus guereza Photo: Anne Savage
	Colobus guereza Photo: Bertrand Deputte
	Colobus guereza Photo: Júlio César Bicca-Marques
	Colobus guereza Photo: Kim Bauers
	Colobus guereza Photo: Kim Bauers
	Colobus guereza Photo: Michael Pogany
	Colobus guereza Photo: Primates in Art & Illustration Collection
	Colobus guereza Photo: Tara Harris

	Colobus guereza occidentalis Photo: Alain Houle
	Colobus guereza occidentalis Photo: Alain Houle
	Colobus guereza occidentalis Photo: Alain Houle
	Colobus guereza occidentalis Photo: Alain Houle

Mantled guereza (colobus)

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ECOLOGY

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INTERNATIONAL STATUS

CONSERVATION THREATS

Threat: Human-Induced Habitat Loss and Degradation

Threat: Invasive Alien Species

Threat: Harvesting (hunting/gathering)

Threat: Persecution

Threat: Changes in Native Species Dynamics

LINKS TO MORE ABOUT CONSERVATION

CONSERVATION INFORMATION

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ORGANIZATIONS INVOLVED IN Colobus guereza CONSERVATION

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