Mangabey SSP

The captive management of mangabeys has been hindered by the lack of information as well as by the difficulty in moving animals (lack of space) to accomplish breeding recommendations.

The climate of Africa has changed numerous times over the last 20,000 years as desert and forest alternately expanded and diminished. Beginning 12,000 to 21,000 years ago, the present day lowland forests were drastically reduced in extent and restricted to small areas in Western and Central Africa (Kingdon, 1981). In these refuge areas, the inhabitants diverged into differing species. When the forests were restored and rejoined (approximately 10,000 years ago), the localized species like Cercocebus and Lophocebus spread (Jolly, 1985).

Thus, the possible historic range for mangabeys stretched from Western to Eastern Africa across the equatorial belt.

In recent history, the forests have not only been reduced by climate changes but have become further isolated and fragmented by exploration (roads, etc.), increased human population and deforestation. In addition to those issues, mangabeys are also hunted as bush meat. Today, mangabey species are found in forest pockets from Guinea, across the continent to Kenya.

Present known ranges for mangabey species in captivity historically in North America (see map):

· Agile Mangabey (C. agilis / C. a. agilis / C. galeritus agilis): Cameroon and Central African Republic, south to Gabon and Democratic Republic of Congo (formerly Zaire).

· Golden-bellied Mangabey (C. chrysogaster/ C. agilis chrysogaster / C. galeritus chrysogaster): Central and Northern Democratic Republic of Congo (DRC), Congo Basin.

· Red-capped Mangabey (C. torquatus / C. t. torquatus): Western and Southern Nigeria, south through Cameroon and Gabon.

· Sooty Mangabey (C. atys atys/ C. torquatus atys): Northern Guinea to Southern Sierra Leone.

· White-crowned Mangabey (C. atys lunulatus/ C. torquatus lunulatus): Eastern Guinea region, Central and Eastern Ivory Coast and Ghana, southern Burkina Faso.

· Grey-cheeked Mangabey (L. albigena): Cameroon to Gabon, northeast Burundi, Rwanda and Uganda.

· Black Mangabey (L. aterrimus / L. albigena aterrimus): Northeastern Angola, DRC.

Present known ranges for mangabey species not in captivity outside of Africa (see map):

· Tana River Mangabey (C. galeritus galeritus): Tana River delta (Southeastern Kenya)

· Sanje Mangabey (C. galeritus sanjei): Udzungwa Mountains (Southwestern Tanzania)

· Highland Mangabey (L. kipunji): Ndundulu Forest of the Udzungwa Mountains and Southern Highlands (Tanzania)

Ecology and Behavior

Not all mangabey species have been studied in the wild. Between 1963 and the early 1980’s, there were studies from the wild dealing largely with population estimates and food and habitat preferences at specific study sites. These studies did not cover all the mangabey species nor were they complete studies of the animals themselves. Fortunately in the last decade, there has been a renewed interest in studying mangabeys in the wild. New information on agile, sooty, white-crowned, Highland and Sanje mangabeys is being published.

The behaviors of Cercocebus and Lophocebus are not well understood nor completely known. Even though mangabeys have been in the North American Region since 1875, little of their complex body posturing or facial displays have been well documented. More importantly, the interesting differences between the species have yet to be fully studied.

All mangabeys appear to have a social organization that can be described as baboon-like. Agonistic interactions between individuals are common and all species are known to have spacing calls that differ greatly in their structure from one another. Life span in captivity commonly reaches the mid to late 20's and the rare individual has made it as far as early 40’s.

Mangabeys are social, medium-sized, African primates that can be separated into two "forms"- the semi-terrestrial "form" such as C. galeritus, C. agilis, C. torquatus and C. atys and the arboreal "form" such as L. albigena and L. aterrimus.

See Table 2 for comparisons and similarities between the genera.

Cercocebus … Semi-terrestrial Mangabey “Forms”

The semi-terrestrial "forms" are markedly sexually dimorphic (females are approximately 20% smaller than males). Head and body length is approximately 22 inches (56 cm) and tail length is approximately 28 inches (72 cm). Mature adult males of C. torquatus, C. agilis and C. galeritus are not only larger in size but very thickset as well, more thickset than mature males of C. atys. The tail is held either parallel to the back (the base forming an acute angle with it) or extended nearly straight behind.

Hands and feet are typical of Old World monkeys with a fully opposable thumb and divergent big toe. Quadrupedal locomotion is digitigrade. Ischial callosities are continuous across the midline on males and separated on females. Large incisor teeth and strong jaws enable them to eat food (i.e. palm nuts) not exploitable by other monkeys such as guenons.

Their feeding time is split between foraging in the trees and foraging through ground leaf-litter. Diet consists mainly of fruits and seeds, with insects forming an important addition (Gautier, Gautier-Hion, 1969; Quris, 1975; Homewood, 1975). Cheek pouches allow food to be gathered for later consumption. Troops may come together briefly to form large groups at plentiful food sources, but in general, they are known to only associate with other primate species haphazardly.

C. galeritus, C. agilis, C. torquatus and C. atys are adapted to swamp forests and other forest types likely to be seasonally or intermittently flooded (Jones, Sabater Pi, 1968; Andrews, Groves, Horne, 1975; Homewood, 1975). C. galeritus sanji is found in much higher elevations, 400 – 1600 m (1300 – 5300 ft), although it is probably more common above 1000 m (3300 ft) (Ehardt, 2001).

Like other semi-terrestrial primates, they generally form multi-male groups ranging in size from 5 to 36 individuals. Because they spend a considerable amount of time on the ground and at lower forest heights (more open) they have developed communication skills such as body postures and facial displays and even possess non-pigmented or white areas above the upper eyelids to enhance the communicative effect of raising the eyebrow. Facial demonstrations such as lip-smacking and open-mouthed threat gestures are common.

Lophocebus … Arboreal Mangabey “Forms”

The arboreal "forms" are only slightly sexually dimorphic (females are approximately 7% smaller than males) with a more slender appearance than the semi-terrestrial "forms" and a longer tail (31 in/ 82 cm) to help with arboreal balance. Nowak (1999) reported tail length in some individuals to reach 100 cm (39 in). When standing still, the tail is generally held at right angle to the back with just the tip forward and when moving, the tail is held behind. They possess an opposable thumb and divergent big toe and like other arboreal primates, quadrupedal locomotion is palmigrade. Ischial callosities are continuous across the midline on males and separated on females. Multi-male group size is smaller (7 to 13) than the semi-terrestrial mangabeys and they come to the ground infrequently.

Diet consists mainly of buds, shoots and fruit. Like other mangabeys they have large cheek pouches and considerable incisors to exploit nuts that other monkeys cannot. These mangabey species are known to form temporary polyspecific associations with other primates and in studies by Gautier, Gautier-Hion (1969) and Gartlan, Struhsaker (1972), they were seen much more commonly in mixed than homogenus troops.

L. albigena and L. aterrimus are a more typical primary and secondary forest primate (Jones, Sabater Pi, 1968) although they do occur in swamp forests (Gautier, Gautier-Hion, 1969; Chalmers, 1968).

The recently discovered L. kipunji lives at elevations up to 2.5 kilometers (1.5 miles) above sea level and endure temperatures as low as -3° Celsius (27° Fahrenheit) and seasonal rainfall that can total almost 3 meters (9.5 feet).

Unlike the semi-terrestrial "forms" which live in a more open habitat, L. albigena and L. aterrimus live in the relatively dense forest canopy. Therefore they lack an extensive repertoire of facial demonstrations since visual displays have a somewhat limited use (they do not have a non-pigmented or white area above the upper eyelids). Instead, they are highly vocal, communicating through a variety of vocalizations such as grunts, calls and barks.

Reproduction

Female mangabeys (as well as baboons and some macaque and guenon species) have perineal swellings on an approximate 30-day cycle. The area around the anus (perineum) increases in size during the first half, or follicular phase of the cycle and decreases after ovulation. Peak swelling is a visual signal advertising ovulation and an optimal time for conception. Menses between cycles is often very visible, especially in the semi-terrestrial mangabeys.

C. torquatus, C. galeritus galeritus, C. chrysogaster and C. atys atys have been documented as exhibiting post-conception perineal swellings. These swellings occur during the last part of the first trimester (the first 1.5 to 2 months of the pregnancy). It is thought that this will to confuse the exact time of conception therefore protecting the infant from possible infanticide from males in the group. Break-through bleeding (most likely from egg implantation) during pregnancy has been documented in C. torquatus and C. chrysogaster.

For some mangabey species there may be some birth seasonality in the wild due to food resources and availability but there does not appear to be any significant birth peaks in captivity. Gestation is approximately 5.5 to 6 months. Twinning has been recorded only once in this region (C. atys atys, 1978), thus twinning is quite rare.

According to the studbook data, females are able to conceive as early as 3.3 years of age and males can sire an offspring as early as 3.5 years of age. These ages, however, are no doubt early compared to the wild.

Male mangabeys become sexually dimorphic at around 5 years of age, so this is probably the normal age of first pregnancy and siring. In captivity, it is important that youngsters have the opportunity to see breeding behavior and the rearing of infants. It is common for immature males to “practice” breeding on different members in the group.

TABLE 2 shows the comparisons and similarities between the two genera

*Cercocebus* Semi-terrestrial Mangabeys	*Lophocebus* Arboreal Mangabeys
Agile mangabey C. agilis / C. a. agilis / C. galeritus agilis Golden-bellied mangabey C. chrysogaster / C. agilis chrysogaster / C. galeritus chrysogaster Red-capped mangabey C. torquatus / C. t. torquatus Sooty mangabey C. atys atys / C. torquatus atys White-crowned mangabey C. atys lunulatus / C. torquatus lunulatus Tana River Mangabey C. galeritus galeritus Sanje Mangabey C. galeritus sanjei	Grey-cheeked mangabey L. albigena Black mangabey L. aterrimus / L. albigena aterrimus Highland Mangabey L. kipunji
Marked sexual dimorphism (females ~ 20% smaller)	Less sexual dimorphism (females ~ 7 % smaller)
Ground foragers, digitigrade locomotion	Infrequently on the ground, palmigrade locomotion
Group size up to 35 with periodic group size reaching 100 individuals (multi-male groups)	Group size often under 20 (single-male or multi-male group composition)
Larger home ranges	Smaller home ranges
Not inclined to form polyspecific associations	Forms occasional polyspecific associations
Various pelage colorations (browns and grays)	A more uniform dark pelage (black hues), expect L. kipunji
Non-pigmented or white patch above the upper lid	No patch above the upper eyelid
Utilize many facial expressions	Utilize many vocalizations
Heavy and visible menses	Menses visible but not as heavy
Natal coat and natal skin differences	Natal skin differences only
Post-conception perineal swellings documented	No post-conception perineal swellings documented
**Cercocebus and Lophocebus What Characteristics They Share**
Estrous cycles (~ 30 days) characterized by perineal swellings
~ 5.5 month gestation
Formidable canine teeth and jaws to access hard nuts (out-competes other primates)
Omnivorous, opportunistic feeders with fruit being an important part of the diet
Ischial callosities and cheek pouches