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Associations

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Sooty mangabey predators include crowned hawk eagles, leopards, humans, and chimpanzees. Sooty mangabeys are noted for their sentinel ability to spot predators on the ground (McGraw and Bshary 2002). Gaboon vipers also elicit alarm calls, although a death from one has yet to be observed in the wild (Range and Fischer 2004). In response to perched African crowned hawk eagles, sooty mangabeys produce an alarm call, and if one is observed flying, the sooty mangabeys descend trees onto the ground. When leopards are observed, sooty mangabeys ascend trees and vocalize alarm calls (Range and Fischer 2004).

Known Predators:

  • chimpanzees (Pan troglodytes)
  • leopards (Panthera pardus)
  • crowned hawk eagles (Stephanoaetus coronatus)
  • humans (Homo sapiens)
  • gaboon vipers (Bitis gabonica)
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Lee, S. 2012. "Cercocebus atys" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Cercocebus_atys.html
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Scarlett Lee, Yale University
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Tanya Dewey, University of Michigan-Ann Arbor
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Morphology

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A member of the Cercopithecinae, monkeys with cheek-pouches, sooty mangabeys are a smoky, slate-grey or brown-grey monkey with lighter white on their ventral side (Groves 2001). They are noted for their light colored facial whiskers, which have the greatest contrast in color in the eastern populations, and hands and feet of slightly darker color than the body. Their bare facial skin is mottled dark grey and pink salmon color, while the upper eyelids are white and the orbits and muzzle are more rectangular than round in outline (Kingdon 1997, Rowe 1996, Meester and Setzer 1971). Their face has a blackish muzzle and their ears are a similar color to their muzzle. Enlargement of the second premolar (P4) relative to the first molar separates the sooty mangabey’s dentition from other closely related species (Daegling et al. 2011). This species is also less sexually dimorphic than Cercocebus torquatus (Groves 2001). The skull of the male is smaller than in C. torquatus and also has a narrower, shorter face. Both sexes also have a significantly lower ascending ramus of the mandible compared to other closely related species (Groves 1978).

Male sooty mangabeys are larger, heavier, and have larger canines than in females (Fruteau, Range, and Noe 2010). The female skull is 87% the size of that of the male (Groves 1978).

The mass of the sooty mangabey ranges from 8.5 to 14 kg for males and 5 to 9 kg for females, with an average of 8.593 kg (Kingdon 1997, Rowe 1996, Mann et al. 1983). The head and body length is 40 to 60 cm in females and 47 to 67 cm in males, while tail length is 40 to 80 cm in both sexes. The height at shoulder for females is 38 to 42 cm and 40 to 45 cm in males (Kingdon 1997).

The skin of the sooty mangabey has unique histological and biochemical properties including subepidermal cholinesterase-reaction nerve fibers over the body surface, and specialized nerve end organs in many areas of the body. There are also specialized nerve end-organs at the base of the epidermal ridges of the fingertips (Machida et al. 1965).

There are two known subspecies of C. atys. The first, Cercocebus atys atys, is noted for being smoky gray, occasionally with a small dorsal stripe but never with a whorl or crown on the head, and crown hairs with a black tip and straw-colored band. The second, Cercocebus atys lunulatus (white collared (crowned) mangabeys) is noted for a whorl or crown on the head, a darker face, a more prominent dorsal stripe, and a white oval mark edged with black on the nape. The hands and feet in white collared mangabeys are more similarly colored to the body and have a white underside (Groves 2001).

Range mass: 5 to 14 kg.

Average mass: 8.6 kg.

Range length: 40 to 67 cm.

Other Physical Features: endothermic ; homoiothermic; bilateral symmetry

Sexual Dimorphism: male larger

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Lee, S. 2012. "Cercocebus atys" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Cercocebus_atys.html
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Scarlett Lee, Yale University
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Rachel Racicot, Yale University
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Tanya Dewey, University of Michigan-Ann Arbor
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Life Expectancy

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Average lifespan in the wild for sooty mangabeys is 18 years (Rowe 1996). Average lifespan in captivity for males is 26.8 years (Carey and Judge 2000).

Average lifespan
Status: wild:
18 years.

Average lifespan
Status: captivity:
26.8 years.

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Lee, S. 2012. "Cercocebus atys" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Cercocebus_atys.html
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Scarlett Lee, Yale University
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Habitat

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Sooty mangabeys are primarily terrestrial and reside in valleys in primary, secondary, flooded, dry, mosaic and mangrove forests in the Guinean Forest Zone. They also inhabit gallery forests and are commonly spotted near swamp and palm forests (Kingdon 1997, Carey and Judge 2000).

The Tai National Park in southwestern Ivory Coast is the only primary forest left in West Africa, and many wild sooty mangabey groups are found and studied extensively there (Range and Noe 2002). In this park, there are two dry seasons and average rainfall is 1,830 mm. The mean temperature is 24 degrees Celsius (McGraw 1998).

Sooty mangabeys live from sea level up to 1000 m above sea leval, possibly higher in the Lome Mountains in Sierra Leone (Oates, Gippoliti, and Groves 2011).

Range elevation: 0 to 1000 m.

Habitat Regions: tropical ; terrestrial

Terrestrial Biomes: forest ; rainforest

Wetlands: swamp

Other Habitat Features: agricultural ; riparian

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Lee, S. 2012. "Cercocebus atys" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Cercocebus_atys.html
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Scarlett Lee, Yale University
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Untitled

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Sooty mangabeys are natural hosts of SIV-2, meaning they acquire the infection but never show any symptoms for it. The captive colony of sooty mangabeys at the Yerkes facility are all infected with SIV-2 and have provided an important study opportunity for HIV researchers. SIV-2 is non-pathogenic in sooty mangabeys (Riddick et al. 2010). One possible route of exposure between these monkeys is through grooming, where blood sucking parasites are removed using the mouth (Santiago et al. 2005). The human genome sequencing center at the Baylor College of Medicine is currently sequencing the genome of sooty mangabeys to understand their “AIDS resistance” factors ("Sooty Mangabey" 2011).

Sooty mangabeys are the only primates other than humans known to acquire leprosy from their own species (Rowe 1996). Leprosy in wild sooty mangabeys, and subsequently in the captive Yerkes sooty mangabey colony, eventually led researchers to discover that Rhesus monkeys infected with SIV were more susceptible to also being infected with leprosy; conversely this led to the discovery that leprosy may prevent AIDS induced mortality (Hamilton et al. 2008).

Sooty mangabeys are often considered a subspecies of Cercocebus torquatus as Cercocebus torquatus atys (Nowak 1991). Cercocebus lunulatus has been proposed as a separate species rather than its current status as a subspecies of Cercocebus atys (Smith 2012).

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Lee, S. 2012. "Cercocebus atys" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Cercocebus_atys.html
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Behavior

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The most common call of sooty mangabeys is a soft grunt, also known as a staccato bark, which consists of a multitude of rapid grunts and a whoop that is finished by a multi-syllabled rumble (Kingdon 1997). Sooty mangabey vocalizations can be grouped into several main categories. Grunts, low frequency vocalizations, are the most common vocalization, and are more commonly produced by males and in foraging activities. Twitters, ranging from soft, melodic sounds to harsher sounds, are only heard from adult females and juveniles of both sexes in foraging and social interactions. They can consist of up to 23 syllables. Twitters can be heard when a female approaches another group member, and are common when foraging in widely dispersed areas to locate another foraging group. Screams, which are loud, noisy agonistic vocalizations, are mostly heard from juvenile and adult females during contact aggression. Growls are multi-syllabic, also occur in agonistic situations, and are often accompanied by a raised eyebrow. Grumbles, hoos, intense threats, and waus are other calls heard during agonistic interactions. Copulation calls have a complex phrase structure and last up to 10 seconds. They can be heard from females during copulation and occasionally during defecation, as well as from males after ejaculation. Whoop gobbles are long calls, only used by adult males, and are associated with contacting other groups. Alarm calls are produced by both males and females, and have been heard in response to Gaboon vipers, leopards, and African crowned eagles. There appears to be a different alarm call specific to each predator (Range and Fischer 2004).

Sooty mangabeys exhibit many variable facial expressions, most of unknown meaning (Kingdon 1997). A lipgrin with a protrusion of the tongue is often seen but its significance is unknown (Bernstein 1976). Frequent eyelid raising indicates a threat (Gust and Gordon 1994).

Communication Channels: visual ; acoustic

Perception Channels: visual ; tactile ; acoustic ; chemical

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Lee, S. 2012. "Cercocebus atys" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Cercocebus_atys.html
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Scarlett Lee, Yale University
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Conservation Status

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While hunting sooty mangabeys is difficult because they live in dense, swampy forests, they are easily trapped since they spend the vast majority of their time foraging on the forest ground (Kingdon 1997). Sooty mangabeys are currently being hunted at two times a sustainable rate in the Tai forest region in Ivory Coast for their meat, which is decimating their population (Refisch and Kone 2005). Human activities and land use also threaten the sooty mangabey, as urbanization has caused deforestation in their natural habitat (Galat and Galat-luong 2006). Large-scale logging has greatly reduced the range of this species (Nowak 1991). For this reason, C. atys is listed as vulnerable according to the IUCN and is in Appendix II of the CITES database (Oates, Gippoliti, and Groves 2011; UNEP-WCMC 2012).

Sooty mangabeys, Cercocebus atys atys, are near threatened, as their population has declined 20 to 25% over the past few decades (Oates, Gippoliti, and Groves 2011). This subspecies is present in two protected areas, Tai National Park and Sapo National Park (Oates, Gippoliti, and Groves 2011).

White-naped mangabeys, Cercocebus atys lunulatus, are endangered, and was one of the 25 most threatened primate species (Galat and Galat-Luong 2006). The population has declined by at least 50% over the past 27 years, largely attributed to habitat destruction and hunting pressure, mostly in Ghana (Oates, Gippoliti, and Groves 2011). The recent addition of the AGEREF/Comoe-Leraba Reserve to the Comoe National park could substantially increase the likelihood of this subspecies’ survival (Galat and Galat-Luong 2006). This park, however, is currently threatened by civil conflict and hunting (Oates, Gippoliti, and Groves 2011). The West African Primate Conservation Action (WAPCA) has highlighted white-naped mangabeys as one of two primate species to save in West Africa and hopes to employ hunters of the monkeys as game wardens to ensure its preservation (Dale 2001).

Captive breeding programs in European zoos also hope to increase population numbers (Oates, Gippoliti, and Groves 2011).

The US Fish & Wildlife Service (USFWS) restrictions currently make continued research on captive sooty mangabeys and SIV challenging. The Yerkes facility is working with collaborators from other institutions to ensure the protection of wild sooty mangabeys, while using colony bred animals to learn more about the basic biology of this species. These institutions hope that their efforts will encourage the USFWS to ease restrictions on the research of the captive colony and SIV (Else et al. 2006).

CITES: appendix ii

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Benefits

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Sooty mangabeys are seen as agricultural pests since they frequently ravage farms, mostly rice paddies, where they can cause extensive economic damage (Kingdon 1997). Attempts to conserve this species may hurt industrialization in the region, since the habitat of this species is composed of trees useful for timber and firewood (Galat and Galat-luong 2006; Oates, Gippoliti, and Groves 2011).

Eastern sooty mangabeys, which are often infected with SIV-2, may have initially been the vectors for SIV-2 to evolve into HIV-2 in humans (Santiago et al. 2005).

Negative Impacts: injures humans (carries human disease); crop pest

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Benefits

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Sooty mangabeys provide a good study primate for evolutionarily, behaviorally, ecologically, and medically related studies. They are studied in the wild at Tai National Park in southwestern Ivory Coast, and in a captive colony at the Yerkes Regional Primate Research Center at Emory University (Fruteau, Range, and Noe 2010; Gust and Gordon 1992). Anthropological research has allowed inferences on human behavior, such as hard object feeding, due to their relatively close phylogenetic relationship to humans (Daegling et al. 2011). This species is also an incredibly important medical study animal as it is a natural host of simian immunodeficiency virus 2 (SIV-2), giving researchers insights on HIV-2 in humans. Sooty mangabeys can also contract and spread leprosy (Hamilton et al. 2008; Riddick et al. 2010).

Sooty mangabeys are hunted for their meat (Refisch and Kone 2005).

Positive Impacts: food ; research and education

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Associations

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Sooty mangabeys follow arboreal monkeys to feed on their dropped fruit. They also act as seed dispersers since their diet includes seeds (Rowe 1996).

Sooty mangabeys have been observed in the wild as participants in mixed primate species groups in the Tai National Park along with western red colobuses (Piliocolobus badius) and Diana monkeys (Cercopithecus diana). These inter-species groups ensure better protection against predators, and usually involve monkeys with different dietary needs to eliminate food competition (Macdonald 2001). These inter-species groups allow arboreal monkeys to come closer to the ground and extend their niche when sooty mangabeys are nearby, due to a perceived reduced threat of terrestrial predators. The sooty mangabey acts as a sentinel for ground predators, and its alarm call helps warn the less observant arboreal species of predators (McGraw and Bshary 2002).

A necroposy of a juvenile female sooty mangabey indicated the presence of lung worms of the Metastrongylidae family, the larval form of Porocephalus armillatus, the pancreatic fluke Brodenia serrata, and Hepatocystis kochi (Hysell et al. 1970). Abbreviata poicilometra, a physalopterid, can also infect this species (Slaughter and Bostrom 1969).

Ecosystem Impact: disperses seeds

Mutualist Species:

  • Diana monkeys (Cercopithecus diana)
  • western red colobus monkeys (Piliocolobus badius)

Commensal/Parasitic Species:

  • Metastrongylidae
  • Porocephalus armillatus
  • Hepatocystis kochi
  • Brodenia serrata
  • Abbreviata poicilometra
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Lee, S. 2012. "Cercocebus atys" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Cercocebus_atys.html
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Trophic Strategy

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Sooty mangabeys are hard object feeders with a narrow diet primarily consisting of fruits and nuts, especially palm nuts, with specialization on their flesh and kernels. They are also known to eat stems, roots, shoots, and leaves of swamp plants, as well as grass, seeds and fungi (Kingdon 1997, Macdonald 2001). Invertebrates are another staple of their diet (McGraw, Vick, and Daegling 2011). They have extremely strong, powerful jaws and massive incisors that allow them to consume a hard object diet. Powerful post-canine biting allows them to crack and consume hard nuts that other animals cannot eat (Rowe 1996, Estes 1991). A complex premolar profile and their thicker enamel also reflect their hard object feeding lifestyle (Daegling et al. 2011).

The most common food in their diet and hardest item they feed on is nuts from Sacoglottis gabonensis, which can comprise 25 to 80% of the whole diet seasonally and which they crush using their large premolars. These nuts are mostly recovered from leaf litter on the forest floor and are covered with an extremely hard casing (McGraw, Vick, and Daegling 2011). When consuming these nuts, they exhibit a powerful isometric bite with incisal preparation and powerful post-canine crushing (McGraw, Vick, and Daegling 2011). The fruit of Anthonota fragrans was observed to seasonally form up to 25% of the diet (Janmaat, Byrne, and Zuberbuhler 2006).

Male and female sooty mangabeys have different dietary patterns. Males tend to eat more hard nuts, seeds, and invertebrates, while females typically consume softer seeds and fruits. Males tend to use more incisions and post-canine crushing. Adults and non-adults also differ in feeding patterns, with adults eating more fungi and soft seeds, and non-adults eating more S. gabonensis seeds (McGraw, Vick, and Daegling 2011).

Animal Foods: insects; terrestrial non-insect arthropods; terrestrial worms

Plant Foods: leaves; roots and tubers; wood, bark, or stems; seeds, grains, and nuts; fruit

Other Foods: fungus

Primary Diet: herbivore (Granivore )

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Lee, S. 2012. "Cercocebus atys" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Cercocebus_atys.html
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Scarlett Lee, Yale University
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Distribution

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Sooty mangabeys are primarily endemic to Upper Guinea in coastal West Africa (Kingdon 1997, Santiago et al. 2005). The native range used to extend from the Casamance River in Senegal to the Sassandra/Nzo River system in Ivory Coast, but their population has been reduced and is mostly extinct today in Senegal, Guinea Bissau, and parts of Guinea (Rowe 1996, Groves 2001). Large numbers of sooty mangabeys still exist in Sierra Leone, Liberia, and western Ivory Coast (Santiago et al. 2005). Their range overlaps with that of other mangabey species (Estes 1991).

Biogeographic Regions: ethiopian (Native )

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Lee, S. 2012. "Cercocebus atys" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Cercocebus_atys.html
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Scarlett Lee, Yale University
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Reproduction

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Sooty mangabeys live in multi-male, multi-female groups of 15 to more than 100 individuals (Stahl and Kaumanns 2003). This social structure has led to a polygynandrous (promiscuous) breeding system, where both males and females have multiple partners (Gouzoules 1984; Fruteau, Range, and Noe 2010).

Although in a promiscuous system females are expected to be aggressive towards each other when competing for mates, no such behavior has been found in the sooty mangabey. However, females are often the victims of aggression from males, and resident males sometimes slap females when they present themselves at the beginning of the breeding season. Males are part of a linear dominance hierarchy, where the highest-ranking male attempts to copulate with all females (Range 2005). Overall, more highly ranked males mount females more often (Fruteau, Range, and Noe 2010).

Non-resident males who visit a group often attack mothers with infants, and often succeed in committing infanticide. Older and higher ranking males defend the females they mated with against such infanticide. After infanticide has been committed, the female immediately goes into estrous, giving the attacking, sneaky male an opportunity to mate with her (Fruteau, Range, and Noe 2010).

Mating System: polygynandrous (promiscuous)

Male sooty mangabeys become sexually active at a little less than 1 year of age, but their earliest ejaculation is at 4 years old. Young males, starting at 1 year, mount sexually mature females. Males actively mount more females from the ages of 3 to 4, during adolescence, than from ages 5 to 6, during sexual maturity (Gust and Gordon 1991). Juvenile males are even more likely to mount sexually mature females than adult males (Rowe 1996). The age of first perineal swelling for females is at 30 to 39 months, around the age of the first menses, and the first birth is usually approximately a year later at 49 to 55 months (Ehardt 1988a, Ehardt 1988b, Mann et al. 1983). The average time between births is 13 to 16 months; having a longer interbirth interval is associated with lower infant mortality rates (Gust and Gordon 1991). Sooty mangabeys give birth to a single infant at a time. Their birthrate is 0.92/yr, and their average gestation period is 167 days (Refisch and Kone 2005).

Female perineal swellings during the estrous cycle last 34.5 days and signal to males that they are capable of mating. Males are able to distinguish between a female’s maximal fertile swelling and a postconception swelling, which occurs during the postconception estrus (Rowe 1996). However, sexual swellings in captive females were not shown to accurately reflect peak fertility, but do represent overall ability to copulate (Whitten and Russell 2006). Females sexually present themselves more frequently to males than to other females, do almost all of the presenting, receive all of the mountings, and perform as many genital inspections as males (Bernstein 1976). More highly ranked males mount females more often than do lower ranked males (Fruteau, Range, and Noe 2010).

Mating first begins with the male grasping the female’s ankles while he mounts her from behind; the female then vocalizes and darts away from him before allowing him to mount her again. Females make a soft grunt during copulation (Gust and Gordon 1994). Females often manually stimulate their perineal area during copulation (Gust and Gordon 1991).

In the wild, mating begins in May, peaks in July to August, and ends in September, although births are sometimes seen throughout the year (Mann et al. 1983). Sooty mangabeys in captivity do not show strong seasonality in mating (Bernstein 1976).

Breeding interval: Breeding occurs once every 13 to 16 months.

Breeding season: The breeding season is from May to September.

Average number of offspring: 1.

Average gestation period: 167 days.

Range weaning age: 4 (low) months.

Average weaning age: 10 months.

Average age at sexual or reproductive maturity (female): 55 months.

Average age at sexual or reproductive maturity (male): 4 years.

Key Reproductive Features: iteroparous ; seasonal breeding ; gonochoric/gonochoristic/dioecious (sexes separate); sexual ; viviparous

At birth, infants are carried on their mother's ventral side but are later carried on the back after a few months. The mother of the infant gives it intensive care for the first 2 to 7 months of life, and more general attention throughout the rest of the first year (Bernstein 1976). The lactation period is 4 to 10 months (Fruteau, Range, and Noe 2010). Aunts of the infant groom the infant from ages 10 to 12 months more than the mother does; siblings, both male and female, also groom infants they are related to (Bernstein 1976). Mothers have also been observed grooming the infants’ eyelashes with a stone (Kyes 1988). Males occasionally carry young to protect the young, but not as a social buffer, since they usually carry infants that they sired themselves (Range and Noe 2002).

Females who lose their infants within six months of their birth immediately re-enter estrus, while mothers who do not lose their infants do not. Therefore, males who kill infants within these six months can reproduce with the female soon after committing infanticide (Fruteau, Range, and Noe 2010).

At birth, the infant is given a rank immediately below the mother, but male offspring quickly rise above the mother in the group ranking after a few years (Bernstein 1976).

Infant neglect and abuse, while not common, has been observed in captive sooty mangabey groups, and is more likely to impact first born young (Maestripieri, Wallen, and Carrol 1997).

Parental Investment: female parental care ; pre-fertilization (Provisioning, Protecting: Male); pre-hatching/birth (Provisioning: Female, Protecting: Female); pre-weaning/fledging (Provisioning: Female, Protecting: Female); pre-independence (Provisioning: Female, Protecting: Female); maternal position in the dominance hierarchy affects status of young

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Lee, S. 2012. "Cercocebus atys" (On-line), Animal Diversity Web. Accessed April 27, 2013 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Cercocebus_atys.html
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Scarlett Lee, Yale University
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Biology

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A rather terrestrial species, the sooty mangabey spends the majority of its time on the forest floor, although it will also climb up into the trees (7). On the forest floor, the sooty mangabey forages primarily for fruit; its large, strong incisor teeth enable it to eat not only the sweet flesh, but also the hard kernels and nuts (4). However, it will also consume many parts of plants found growing in swamps and the seeds of grasses, and will frequently raid farms, including rice paddies, where it feeds on crops causing significant damage (2). Mangabeys typically live in large groups, each containing several adult males (4). In captivity, young were born at any time of the year, but births were more common between October and March (9), after a gestation period of around six months (4).
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Conservation

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The sooty mangabey occurs in a number of protected areas, such as Taï National Park in Côte d'Ivoire (6), as does the white-collared mangabey, which can be found in several parks and reserves including Yoyo Forest Reserve in Ghana and the Marahoué National Park in Côte d'Ivoire (7). To protect this Critically Endangered subspecies from extinction, it has been recommended that those forests containing the remaining populations should be elevated to National Park status, in combination with the instigation of regular patrols to deter hunters (7). Sadly, a lack of local support is said to hinder conservation efforts, and recent civil conflict adds to the difficulties in carrying out conservation work in this region (7). In 2001, a group of European zoos decided to collaborate under the name of West African Primate Conservation Action (WAPCA), with the aim of securing the future of both the white-collared mangabey and the Roloway monkey (Ceropithecus diana roloway) which inhabits the same region. WAPCA runs a centre in Ghana, which helps breed both monkeys as part of an international breeding programme, in addition to undertaking research and raising awareness in local communities (10).
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Description

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Mangabeys are medium-sized monkeys of Africa, all noted for their tails which are longer than their bodies (4). In keeping with its name, the sooty mangabey has smoky-grey fur with lighter underparts and slightly darker hands and feet (2). It has a fairly long muzzle bearing whiskers (2) (5), and the bare facial skin is dark grey or mottled, except for the upper eyelids which are strikingly white. Flashes of these conspicuous eyelids, in combination with a variety of facial expressions, are used to communicate (2). Two subspecies are recognised: Cercocebus atys atys and Cercocebus atys lunulatus, the white-collared mangabey. As its name suggests, the white-collared mangabey has a prominent collar of white fur (2). More often heard than see (2), the sooty mangabey has a rather broad vocal repertoire, including grunts and twitters which are frequently heard when foraging or during social interactions, and screams and growls given during aggressive and defensive situations (6).
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Habitat

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This monkey occurs in a broad range of habitats, including primary and secondary forests (2), swamp and agricultural areas (7).
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Range

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The sooty mangabey occurs in West Africa. C. a. atys is found in southern Guinea, east to the River Sassandra in Côte d'Ivoire, while the Critically Endangered white-collared mangabey (C. a. lunulatus) occurs in just a few remaining forest patches between the River Sassandra, Côte d'Ivoire and the River Volta, Ghana (2) (7), and in Burkina Faso (8).
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Status

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Classified as Vulnerable (VU) on the IUCN Red List (1) and listed on Appendix II of CITES (3). Subspecies: Cercocebus atys atys is classified as Near Threatened (NT) and Cercocebus atys lunulatus (the white-collared mangabey or white-naped mangabey) is classified as Endangered (EN) on the IUCN Red List (1).
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Threats

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Whilst this species, like many other primates, is impacted by both hunting and habitat loss (2), it is still widespread and is therefore not currently considered to be threatened with extinction (1). However, the outlook for the white-collared mangabey (subspecies C. a. lunulatus), is far grimmer, as hunting and habitat loss has forced this monkey into just a few remaining patches of forest, and to the brink of extinction (7). Forests in parts of the sooty mangabey's range have been reduced to less than ten percent of their original size, and as the forests become smaller and more fragmented, previously inaccessible areas are opened up, increasing hunting pressure on the remaining monkey populations (7). Unfortunately, the ground-foraging habits of the sooty mangabey make it highly susceptible to being caught in traps laid on the forest floor, and in many agricultural areas, this primate is treated as a pest due to the damage it can do to crops (2).
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Sooty mangabey

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The sooty mangabey (Cercocebus atys) is an Old World monkey found in forests from Senegal in a margin along the coast down to the Ivory Coast.[1]

Habitat and ecology

The sooty mangabey is native to tropical West Africa, being found in Guinea, Guinea-Bissau, Liberia, Senegal, Sierra Leone, and Ivory Coast.[2] Sooty mangabeys inhabit both old growth and secondary forests as well as in flooded, dry, swamp, mangrove, and gallery forests. Sooty mangabeys are terrestrial omnivores, typically spending around 75% of their overall time on the ground (~85% of travel time and ~71% of foraging time).[3] In their foraging behaviors, sooty mangabeys typically consume fruits (~20% of diet), invertebrates (~13% of diet), and nuts and seeds (>55% of diet).[4] In acquiring nuts, sooty mangabeys have been observed scavenging the remains of coula and panda nuts cracked by chimpanzees and red river hogs, potentially using either the sound of cracking nuts or social networks to identify sites of remnants.[5]

Taxonomy

Until 2016, Cecrocebus atys was considered a single species with two subspecies of this mangabey: Cecrocebus atys atys (now Cercocebus atys) and Cecrocebus atys lunulatus.[6] After assessment by the International Union for Conservation of Nature (IUCN) in 2016, Cercocebus atys lunulatus was declared a separate species (Cercocebus lunulatus).[7] Both Cercocebus atys and Cercocebus lunulatus were formerly considered subspecies of the widespread Cercocebus torquatus.[1]

  • Cercocebus atys atys, now Cercocebus atys, is commonly known as the sooty mangabey and is situated west of the Sassandra River in the Ivory Coast up to Senegal.[2]
  • Cercocebus atys lunulatus, now Cercocebus lunulatus, is commonly known as the white-crowned,[8] white-naped,[9] or white-collared mangabey (leading to confusion with the collared mangabey).[7] This now distinct species is considered to have a geographic range east of the Sassandra River to the west of the Volta River in Ghana; Cercocebus lunulatus is also recorded as inhabiting forests in southwestern Burkina Faso and northeastern Ivory Coast. Cercocebus lunulatus is considered Endangered by the IUCN.[7]

Appearance

Sooty mangabeys are gray-colored primates with a lighter-colored chest and stomach. Their faces are typically grayish pink, with darker fur along the forehead and ears;[1] given their diet of hard seeds and nuts, sooty mangabeys are observed to have strong molars.[10][4] Sooty mangabeys also show sexual dimorphism; males typically weigh about 10–11 kg (22–24 lb), while females are typically smaller at about 5–6 kg (11–13 lb).[4]

Behavior

Social organization

Sooty mangabeys typically live and forage in large, multi-male, multi-female groups of 70–120 individuals.[11][12] Sooty mangabeys form linear dominance hierarchies within sexes and form coalitions; within these hierarchies, higher-ranking females typically are found to spend less time foraging as opposed to feeding than their lower-ranking counterparts and were more centrally located within groups.[13][14] Similarly, higher-ranking males were found to be more centrally located within the group, and be better fed and rested.[14] And in captivity, higher-ranking males sired more offspring, indicating that higher male rank is generally predictive of greater reproductive success.[15] Overall, however, females are found to be located in a more central spatial position within the group and better fed and rested than males, independent of ranking.[14]

Dominance rankings are not static; turnover of the dominant, alpha male has been recorded.[16] Furthermore, the dominance rank of children is not influenced by the dominance ranking of either parent, and juveniles typically challenge higher ranking adults starting around three or four years of age.[17] Typically, males will outrank all of the females by age five or six.[17]

Communication

Sooty mangabeys are typically predated upon by leopards, eagles, chimpanzees, vipers, and humans.[18] As a result of these selective pressures, sooty mangabeys have evolved acoustically distinct alarm calls for different predator types.[12] These calls are not vocalized specifically in favor of kin or cooperation partners[19] and in fact are used by other monkey species to avoid potential predators.[20]

Sooty mangabeys also produce other vocalizations within their varied repertoire for a wide variety of social interactions.[21] Sooty mangabeys are recorded most frequently producing grunts (typically in the context of foraging, socially embracing, or, between males, for asserting dominance), twitters (typically produced by adult females during foraging and social interactions such as grooming), and screams (emitted during agonistic interactions, typically by juveniles and adult females).[18] Other notable vocalizations include copulation calls mainly emitted by females during intercourse and "whoop gobbles"—low frequency, extended calls emitted by males at a high volume during the morning, with a nearby group, or with sightings or attacks of predators.[18]

When approaching other females with infants, females will use grunts and twitters to signal benign intent.[22] This often leads to unreciprocated grooming from the approaching female—mothers, upon receiving grooming, will allow for the grooming female to handle their infants.[23]

Sexual and reproductive behavior

Female sooty mangabeys have sexual swellings that are maximally tumescent near ovulation and typically have a gestation length of ~160–170 days;[24] while typically, higher ranking males would be able to identify estrous females and monopolize mating opportunities, it is suggested that dominant males cannot entirely control access to estrous females,[15] perhaps because swellings allow females to precipitate paternity confusion through polygynandry.[25] However, despite these potential counterstrategies against infanticide through paternity confusion, cases of infanticide have been recorded, usually shortly after a change in alpha males or with the introduction of new, immigrant males.[26][27]

In captivity, recently deposed alpha males have been observed carrying their infants (likely for protection) in the presence of newly ascended alpha males, typically following aggression by the new alpha male towards the infant.[16] In habituated sooty mangabeys, immigrant males new to the group have been found to attack infants, who would be defend by their mothers.[27] In this context, resident adult males who had mated with the mother (and potentially fathered the infant) were found to defend the mother and infant from the attacking immigrant male.[27]

Females have thus developed behavioral counter-strategies to protect against attacks and infanticide. Females were found to mate with resident males during previous mating seasons and remain in close proximity to these resident males after birth.[27] In addition, females have been found to respond differently to the vocalizations of members of their own group (as opposed to non-group members), suggesting an ability to recognize infanticide threats from strangers.[28]

Disease

Sooty mangabeys are naturally infected with a strain of Simian Immunodeficiency Virus (SIV), known as SIVsmm. Due to extensive human-mangabey contact in sub-Saharan Africa, SIVsmm has jumped from this species into humans on many occasions, resulting in HIV-2 virus.[29][30] Because sooty mangabeys, as natural hosts of SIV, do not get sick from SIV, much research has been performed on the species for potential genetic resistance or immunological mechanisms.[31] The HIV-1 strain by contrast came from the common chimpanzee strain of SIV.[32][33]

Sooty mangabeys can also contract leprosy, caused by the bacterium Mycobacterium leprae.[34] It is one of several species in which naturally acquired leprosy has been reported, the others being humans, the nine-banded armadillo, the common chimpanzee, and the crab-eating macaque; murine leprosy has also been reported in rats and mice, caused by Mycobacerium lepraemurium.[34]

Conservation status

The sooty mangabey is believed to be decreasing in numbers as its forest habitat is degraded, with trees being felled for firewood and timber and forest habitats used for agriculture.[35] Furthermore, sooty mangabeys are hunted for meat in some parts of its range, often at rates far exceeding the rate at which Sooty mangabeys can reproductively sustain themselves; this increase in hunting, especially with improved technology and an influx of human populations (and thus hunters), has become an increasing threat to the conservation of sooty mangabeys.[36] The International Union for Conservation of Nature has assessed the conservation status of sooty mangabeys as Vulnerable.[2]

References

  1. ^ a b c d Groves, C. P. (2005). Wilson, D. E.; Reeder, D. M. (eds.). Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Baltimore: Johns Hopkins University Press. p. 153. ISBN 0-801-88221-4. OCLC 62265494.
  2. ^ a b c d Koné, I.; McGraw, S.; Gonedelé Bi, S.; Barrie, A. (2020). "Cercocebus atys". IUCN Red List of Threatened Species. 2020: e.T136933A92248451. doi:10.2305/IUCN.UK.2020-2.RLTS.T136933A92248451.en. Retrieved 19 November 2021.
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  20. ^ McGraw, W. Scott; Bshary, Redouan (2002). "Association of Terrestrial Mangabeys (Cercocebus atys) with Arboreal Monkeys: Experimental Evidence for the Effects of Reduced Ground Predator Pressure on Habitat Use". International Journal of Primatology. 23 (2): 311–325. doi:10.1023/A:1013883528244. S2CID 37003870.
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  22. ^ Fedurek, Pawel; Neumann, Christof; Bouquet, Yaëlle; Mercier, Stéphanie; Magris, Martina; Quintero, Fredy; Zuberbühler, Klaus (2019). "Behavioural patterns of vocal greeting production in four primate species". Royal Society Open Science. 6 (4): 182181. Bibcode:2019RSOS....682181F. doi:10.1098/rsos.182181. PMC 6502363. PMID 31183141.
  23. ^ Fruteau, Cécile; van de Waal, Erica; van Damme, Eric; Noë, Ronald (2011). "Infant access and handling in sooty mangabeys and vervet monkeys". Animal Behaviour. 81 (1): 153–161. doi:10.1016/j.anbehav.2010.09.028. S2CID 53180842.
  24. ^ Stevenson, Miranda (1973). "Notes on pregnancy in the Sooty mangabey Cercocebus atys". International Zoo Yearbook. 13 (1): 134–135. doi:10.1111/j.1748-1090.1973.tb02126.x.
  25. ^ Alberts, Susan C.; Fitzpatrick, Courtney L. (2012). "Paternal care and the evolution of exaggerated sexual swellings in primates". Behavioral Ecology. 23 (4): 699–706. doi:10.1093/beheco/ars052. PMC 3999376. PMID 24771988.
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  27. ^ a b c d Fruteau, Cécile; Range, Friederike; Noë, Ronald (2010). "Infanticide risk and infant defence in multi-male free-ranging sooty mangabeys, Cercocebus atys". Behavioural Processes. 83 (1): 113–118. doi:10.1016/j.beproc.2009.11.004. PMID 19914358. S2CID 22015547.
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  30. ^ Ayouba, Ahidjo; Akoua-Koffi, Chantal; Calvignac-Spencer, Sébastien; Esteban, Amandine; Locatelli, Sabrina; Li, Hui; Li, Yingying; Hahn, Beatrice H.; Delaporte, Eric; Leendertz, Fabian H.; Peeters, Martine (2013-09-24). "Evidence for continuing cross-species transmission of SIVsmm to humans: characterization of a new HIV-2 lineage in rural Côte d'Ivoire". AIDS. 27 (15): 2488–2491. doi:10.1097/01.aids.0000432443.22684.50. PMC 3881176. PMID 23939239.
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  34. ^ a b Rojas-Espinosa O, Løvik M (2001). "Mycobacterium leprae and Mycobacterium lepraemurium infections in domestic and wild animals". Rev. Sci. Tech. Off. Int. Epiz. 20 (1): 219–51. doi:10.20506/rst.20.1.1271. PMID 11288514.
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Sooty mangabey: Brief Summary

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The sooty mangabey (Cercocebus atys) is an Old World monkey found in forests from Senegal in a margin along the coast down to the Ivory Coast.

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