Gelada: Difference between revisions

Gelada
	Male
	Female with baby drinking; Both near Debre Libanos, Ethiopia
Conservation status
	; Least Concern (IUCN 3.1)
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Order:	Primates
Suborder:	Haplorhini
Infraorder:	Simiiformes
Family:	Cercopithecidae
Subfamily:	Cercopithecinae
Tribe:	Papionini
Genus:	Theropithecus; I. Geoffroy, 1843
Species:	T. gelada
Binomial name
	Theropithecus gelada; (Rüppell, 1835)
	Gelada range

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The gelada (Theropithecus gelada, Template:Lang-am), sometimes called the bleeding-heart monkey or the gelada "baboon", is a species of Old World monkey found only in the Ethiopian Highlands, with large populations in the Semien Mountains. Geladas are actually not baboons (baboons are all taxonomic members of the genus Papio) but members of their own genus Theropithecus. Theropithecus is derived from the Greek root words for "beast-ape".^[3]^[4] Like its close relatives the baboons (genus Papio), it is largely terrestrial, spending much of its time foraging in grasslands.

Phylogeny and fossils

Since 1979, it has been customary to place the gelada in its own genus (Theropithecus), though some genetic research suggests that this monkey should be grouped with its papionine (baboon) kin;^[5] other researchers have classified the species even farther distant from Papio.^[6] While Theropithecus gelada is the only living species of its genus, separate, larger species are known from the fossil record: T. brumpti, T. darti^[7] and T. oswaldi, formerly classified under genus Simopithecus.^[8] Theropithecus, while restricted at present to Ethiopia, is also known from fossil specimens found in Africa and the Mediterranean into Asia, including South Africa, Malawi, the Democratic Republic of the Congo, Tanzania, Uganda, Kenya, Ethiopia, Algeria, Morocco, Spain, and India, more exactly at Mirzapur, Cueva Victoria, Pirro Nord, Ternifine, Hadar, Turkana, Makapansgat and Swartkrans.

The two subspecies of gelada are:^[2]

Northern gelada, Theropithecus gelada gelada
Eastern gelada, southern gelada or Heuglin's gelada, Theropithecus gelada obscurus

Physical description

The gelada is large and robust. It is covered with buff to dark brown, coarse hair and has a dark face with pale eyelids. Its arms and feet are nearly black. Its short tail ends in a tuft of hair.^[9]^[10] Adult males have a long, heavy cape of hair on their backs.^[9]^[10] The gelada has a hairless face with a short muzzle that is closer to a chimpanzee's than a baboon's.^[10] It can also be physically distinguished from a baboon by the bright patch of skin on its chest.^[9]^[10] This patch is hourglass-shaped. On males, it is bright red and surrounded by white hair; on females, it is far less pronounced. However, when in estrus, the female's patch will brighten, and a "necklace" of fluid-filled blisters forms on the patch. This is thought to be analogous to the swollen buttocks common to most baboons experiencing estrus. In addition, females have knobs of skin around their patches. Geladas also have well developed ischial callosities.^[10] There is sexual dimorphism in this species: males average 18.5 kg (40.8 lb) while females are smaller, averaging 11 kg (24.3 lb).^[11] The head and body length of this species is 50–75 cm (19.7–29.5 in) for both sexes. Tail length is 30–50 cm (11.8–19.7 in).^[10]

The gelada has several adaptations for its terrestrial and graminivorous (grass-eating) lifestyle. It has small, sturdy fingers adapted for pulling grass and narrow, small incisors adapted for chewing it. The gelada has a unique gait, known as the shuffle gait, that it uses when feeding.^[12] It squats bipedally and moves by sliding its feet without changing its posture.^[12] Because of this gait, the gelada's rump is hidden beneath and so unavailable for display; its bright red chest patch is visible, though.

male
female
male head
juvenile male
female with baby on her back

Range and ecology

Grazing geladas, at 3,000 m (9,800 ft) in the Semien Mountains

Geladas are found only in the high grassland of the deep gorges of the central Ethiopian plateau. They live in elevations 1,800–4,400 m (5,900–14,400 ft) above sea level, using the cliffs for sleeping and montane grasslands for foraging. These grasslands have widely spaced trees and also contain bushes and dense thickets.^[9]^[13] The highland areas where they live tend to be cooler and less arid than lowlands areas.^[13] Thus, the geladas usually do not experience the negative effects that the dry season has on food availability. Nevertheless, in some areas, they do experience frost in the dry season, as well as hailstorms in the wet season.

Geladas are the only primates that are primarily graminivores and grazers – grass blades make up to 90% of their diet. They eat both the blades and the seeds of grasses. When both blades and seeds are available, geladas prefer the seeds. They eat flowers, rhizomes and roots when available,^[12]^[13] using their hands to dig for the latter two. They consume herbs, small plants, fruits, creepers, bushes and thistles.^[12]^[13] Insects can be eaten, but only rarely and only if they can easily be obtained. During the dry season, grasses are eaten less and herbs are preferred. Geladas consume their food more like ungulates than primates, and they can chew their food as effectively as zebra.^[14]

Geladas are primarily diurnal. At night, they sleep on the ledges of cliffs.^[15] At sunrise, they leave the cliffs and travel to the tops of the plateaux to feed and socialize.^[12] When morning ends, social activities tend to wane and the geladas primarily focus on foraging. They will travel during this time, as well. When evening arrives, geladas exhibit more social activities before descending to the cliffs to sleep.^[12]

Behavior

Social structure

Geladas live in a complex multilevel society similar to that of the hamadryas baboon. The smallest and most basic groups are the reproductive units, which are made up of one to twelve females, their young and one to four males, and the all-male units, which are made up of two to fifteen males. The next level of gelada societies are the bands, which are made up of two to 27 reproductive units and several all-male units. Herds consist of up to 60 reproductive units that are sometimes from different bands and last for short periods of time. Communities are made of one to four bands whose home ranges overlap extensively. A gelada can typically live to around 20 years old.^[15]^[16]^[17]

Within the reproductive units, the females tend to be closely related and have strong social bonds.^[16] Reproductive units split if they become too large. While females have strong social bonds in the group, a female will only interact with at most three other members of her unit.^[16] Grooming and other social interactions among females usually occur between pairs.^[18] Females in a reproductive unit exist in a hierarchy, with higher-ranking females having more reproductive success and more offspring than lower-ranking females.^[19] Closely related females tend to have a similar hierarchical status.^[19] Females generally stay in their natal units for life; cases of females leaving are rare.^[20] Aggression within a reproduction unit, which is rare, is usually just between the females.^[18] Aggression is more frequent between members of different reproductive units and is usually started by females, but males and females from both sides will join and engage if the conflict escalates.^[18]

Males can remain in a reproductive unit for four to five years.^[16] While geladas have traditionally been considered to have a male-transfer society, many males appear to be likely to return and breed in their natal bands. Nevertheless, gelada males leave their natal units and try to take over a unit of their own. A male can take over a reproductive unit either through direct aggression and fighting or by joining one as a subordinate and taking some females with him to create a new unit.^[16] When more than one male is in a unit, only one of them can mate with the females.^[18]^[20] The females in the group together can have power over the dominant male. When a new male tries to take over a unit and overthrow the resident male, the females can choose to support or oppose him. The male maintains his relationship with the females by grooming them rather than forcing his dominance, in contrast to the society of the hamadryas baboon. Females accept a male into the unit by presenting themselves to him. Not all the females may interact with the male. Usually, one may serve as his main partner.^[21] The male may sometimes be monopolized by this female.^[21] The male may try to interact with the other females, but they are usually unresponsive.^[21]

Most all-male units consist of several subadults and one young adult, led by one male. A member of an all-male unit may spend two to four years in the group before attempting to join a reproductive unit. All-male groups are generally aggressive towards both reproductive units and other all-male units.^[18] As in reproductive units, aggression within all-male units is rare. As bands, reproductive units exist in a common home range.^[22] Within the band, members are closely related and between the units there is no social hierarchy. Bands usually break apart every eight to nine years as a new band forms in a new home range.

Researchers from the University of the Free State (UFS) in South Africa, while observing gelada during field studies, discovered that the monkeys were capable of 'cheating' on their partners and covering up their 'infidelity'. A non-dominant male would mate surreptitiously with a female, suppressing their normal mating cries so as not to be overheard. If discovered, the dominant male would attack the miscreants in a clear form of punishment. It is the first time that evidence of the knowledge of cheating and fear of discovery has been recorded among animals in the wild. Dr. Aliza le Roux of the university's Department of Zoology and Entomology believes that dishonesty and punishment are not uniquely human traits, and that the observed evidence of this behaviour among gelada monkeys suggests that the roots of the human system of deceit, crime and punishment lie very deep indeed.^[23]

Mixed-species association was observed between solitary Ethiopian wolves and geladas. According to the study's findings, gelada monkeys would not typically move on encountering Ethiopian wolves, even when they were in the middle of the herd—68 percent of encounters resulted in no movement and only 11 percent resulted in a movement of greater than 10 m (33 ft). In stark contrast, the geladas always fled great distances to the cliffs for safety whenever they encountered aggressive domestic dogs.^[24]

Reproduction and parenting

When in estrus, the female points her posterior towards a male and raises it, moving her tail to one side.^[25] The male then approaches the female and inspects her chest and genital areas.^[25]^[26] A female will copulate up to five times per day, usually around midday.^[26] Breeding and reproduction can occur at any time of the year, although some areas have birth peaks.^[11]^[27]

Gelada displaying its teeth and gums with its lip flipped back

Most births occur at night. Newborn infants have red faces and closed eyes, and they are covered in black hair.^[26] On average, newborn infants weigh 464 g (16.4 oz).^[28] Females that have just given birth stay on the periphery of the reproductive unit. Other adult females may take an interest in the infants and even kidnap them.^[26] An infant is carried on its mother’s belly for the first five weeks, and thereafter on her back.^[26]^[29] Infants can move independently at around five months old. A subordinate male in a reproductive unit may help care for an infant when it is six months old.^[26] When herds form, juveniles and infants may gather into play groups of around ten individuals. When males reach puberty, they gather into unstable groups independent of the reproductive units. Females sexually mature at around three years, but do not give birth for another year.^[18]^[22] Males reach puberty at about four or five years, but they are usually unable to reproduce because of social constraints and wait until they are about eight to ten years old.^[11] Average life span in the wild is 19 years.^[30]

Communication

Adult geladas use a diverse repertoire of vocalizations for various purposes, such as: contact, reassurance, appeasement, solicitation, ambivalence, aggression and defense.^[31] The level of complexity of these vocalizations is thought to near that of humans.^[32] They sit around and chatter at each other, signifying to those around that they matter, in a way, to the individual "speaking". To some extent, calls are related to the status of an individual. In addition, females have calls signaling their estrus. Geladas communicate through gestures, as well. They display threats by flipping their upper lips back on their nostrils to display their teeth and gums, and by pulling back their scalps to display the pale eyelids.^[33] A gelada submits by fleeing or presenting itself.

Conservation status

In 2008, the IUCN assessed the gelada as Least Concern, although their population had reduced from an estimated 440,000 in the 1970s to around 200,000 in 2008. It is listed in Appendix II of CITES.^[2] Major threats to the gelada are a reduction of their range as a result of agricultural expansion and shooting as crop pests. Previously, these monkeys were trapped for use as laboratory animals or hunted to obtain their capes to make items of clothing.^[2] As of 2008, proposals have been made for a new Blue Nile Gorges National Park and Indeltu (Shebelle) Gorges Reserve to protect larger numbers.^[2]

References

^ 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. 167. ISBN 0-801-88221-4. OCLC 62265494.
^ ^a ^b ^c ^d ^e Gippoliti, S.; Hunter, C. (2008). "Theropithecus gelada". The IUCN Red List of Threatened Species. 2008: e.T21744A9316114. doi:10.2305/IUCN.UK.2008.RLTS.T21744A9316114.en. Retrieved 12 January 2018. {{cite journal}}: Unknown parameter |last-author-amp= ignored (|name-list-style= suggested) (help)
^ "Classic Roots P". PHTHIRAPTERA CENTRAL. Retrieved 2006-12-26. thero (G) - A wild beast; summer; hunt for
^ "Classic Roots T". PHTHIRAPTERA CENTRAL. Retrieved 2006-12-26. pithec, -o, -us (G) - An ape
^ Goodman, M.; Porter, C. A.; Czelusniak, J.; Page, S. L.; Schneider, H.; Shoshani, J.; Gunnell, G.; Groves, C. P. (1998). "Toward a Phylogenetic Classification of Primates Based on DNA Evidence Complemented by Fossil Evidence". Molecular Phylogenetics and Evolution. 9 (3): 585–598. doi:10.1006/mpev.1998.0495. PMID 9668008.
^ McKenna, M.C.; Bell, S.K. (1997). Classification of mammals above the species level. New York: Columbia University Press. pp. 631 pp.
^ Hughes, K.; Elton, S.; O'Regan, J. (Jan 2008). "Theropithecus and 'Out of Africa' dispersal in the Plio-Pleistocene". Journal of Human Evolution. 54 (1): 43–77. doi:10.1016/j.jhevol.2007.06.004. ISSN 0047-2484. PMID 17868778.
^ Maier, W. (1972). "The first complete skull of Simopithecus darti from Makapansgat, South Africa, and its systematic position". Journal of Human Evolution. 1 (4): 395–400. doi:10.1016/0047-2484(72)90116-9.
^ ^a ^b ^c ^d Napier PH. (1981). Catalogue of primates in the British museum (natural history) and elsewhere in the British Isles, part II: family Cercopithecidae, subfamily Cercopithecinae, London: British Museum (Natural History).
^ ^a ^b ^c ^d ^e ^f Ankel-Simons F. (2007). Primate Anatomy: an introduction, (3rd edition), San Diego: Elsevier Academic Press.
^ ^a ^b ^c Jolly CJ. (2007). "Baboons, mandrills, and mangabeys: afro-papionin socioecology in a phylogenetic perspective", In: Primates in perspective, Campbell CJ, Fuentes A, MacKinnon KC, Panger M, Bearder SK (eds), New York: Oxford U Pr. p 240-51.
^ ^a ^b ^c ^d ^e ^f Dunbar RIM. (1977). "Feeding ecology of gelada baboons: a preliminary report", In: Primate ecology: studies of feeding and ranging behaviour in lemurs, monkeys and apes, Clutton-Brock TH (ed), London: Academic Pr. p 251-73.
^ ^a ^b ^c ^d Iwamoto T, Dunbar RIM. (1983). "Thermoregulation, habitat quality and the behavioural ecology of gelada baboons", J Anim Ecol 52(2):357-66.
^ Iwamoto, T. (1979). "Ecological and sociological studies of gelada. Feeding ecology". Contributions to Primatology. 16: 279–330. PMID 101341.
^ ^a ^b Crook JH. (1966). "Gelada baboon herd structure and movement: a comparative report", Symp Zool Soc Lond 18:237-58.
^ ^a ^b ^c ^d ^e Dunbar RIM. (1986). "The social ecology of gelada baboons", In: Ecological aspects of social evolution: birds and mammals, Rubenstein DI, Wrangham RW, (eds), Primceton (NJ):Princeton U Pr. pp.332-51.
^ Grüter CC, Zinner D. (2004). "Nested societies. Convergent adaptations of baboons and snub-nosed monkeys?" Prim Rep 70:1-98.
^ ^a ^b ^c ^d ^e ^f Dunbar, R.; Dunbar, P. (1975). "Social dynamics of gelada baboons". Contributions to Primatology. 6: 1–157. PMID 808371.
^ ^a ^b Dunbar, R. I. M. (1980). "Determinants and evolutionary consequences of dominance among female gelada baboons". Behavioral Ecology and Sociobiology. 7 (4): 253–265. doi:10.1007/BF00300665.
^ ^a ^b Ohsawa, H. (1979). "The local gelada population and environment of the Gich area". Contributions to Primatology. 16: 3–45. PMID 101342.
^ ^a ^b ^c Dunbar, R. I. M. (1983). "Structure of Gelada Baboon Reproductive Units III. The Males Relationship with his Females", Animal Behavior 31:565-575.
^ ^a ^b Dunbar, R. I. M. (1984). Reproductive decisions: an economic analysis of gelada baboon social strategies. Princeton, N.J: Princeton University Press. ISBN 978-0-691-08360-5.
^ le Roux, Aliza, Report, Department of Zoology and Entomology, Qwa Qwa campus, University of the Free State {{citation}}: Missing or empty |title= (help)
^ "Wolves are better hunters when monkeys are around: An unexpected co-existence in the Ethiopian highlands". Science Daily. 22 June 2015.
^ ^a ^b Bernstein, I. S. (1975). "Activity patterns in a gelada monkey group". Folia Primatologica. 23 (1–2): 50–71. doi:10.1159/000155661. PMID 806510.
^ ^a ^b ^c ^d ^e ^f Mori, U. (1979). "Ecological and sociological studies of gelada baboons. Reproductive behavior". Contributions to Primatology. 16: 183–197. PMID 101336.
^ Dunbar, R. I. M.; Hannah-Stewart, L.; Dunbar, P. (2002). "Forage quality and the costs of lactation for female gelada baboons". Animal Behaviour. 64 (5): 801. doi:10.1006/anbe.2002.9972.
^ Leutenegger, W. (1973). "Maternal-fetal weight relationships in primates". Folia Primatologica. 20 (4): 280–293. doi:10.1159/000155580. PMID 4208250.
^ Barrett, L.; Dunbar, R. I. M.; Dunbar, P. (1995). "Mother-infant contact as contingent behaviour in gelada baboons". Animal Behaviour. 49 (3): 805. doi:10.1016/0003-3472(95)80211-8.
^ "Gelada". nationalgeographic.com. National Geographic Society. 2011-05-10T06:34:00-0400. Retrieved 7 March 2012. {{cite web}}: Check date values in: |date= (help)
^ Kawai, M. (1979). "Ecological and sociological studies of gelada baboons. Auditory communication and social relations". Contributions to Primatology. 16: 219–241. PMID 101338.
^ Richman, Bruce (1976). "Some vocal distinctive features used by gelada monkeys". Journal of the Acoustical Society of America. 60 (3): 718–724. Bibcode:1976ASAJ...60..718R. doi:10.1121/1.381144. PMID 824335.
^ Mori, U. (1979). "Ecological and sociological studies of gelada baboons. Individual relationships within a unit". Contributions to Primatology. 16: 93–124. PMID 101345.

External links

[MSW3-1] 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. 167. ISBN 0-801-88221-4. OCLC 62265494.

[iucn-2] Gippoliti, S.; Hunter, C. (2008). "Theropithecus gelada". The IUCN Red List of Threatened Species. 2008: e.T21744A9316114. doi:10.2305/IUCN.UK.2008.RLTS.T21744A9316114.en. Retrieved 12 January 2018. {{cite journal}}: Unknown parameter |last-author-amp= ignored (|name-list-style= suggested) (help)

[3] "Classic Roots P". PHTHIRAPTERA CENTRAL. Retrieved 2006-12-26. thero (G) - A wild beast; summer; hunt for

[4] "Classic Roots T". PHTHIRAPTERA CENTRAL. Retrieved 2006-12-26. pithec, -o, -us (G) - An ape

[Goodman-5] Goodman, M.; Porter, C. A.; Czelusniak, J.; Page, S. L.; Schneider, H.; Shoshani, J.; Gunnell, G.; Groves, C. P. (1998). "Toward a Phylogenetic Classification of Primates Based on DNA Evidence Complemented by Fossil Evidence". Molecular Phylogenetics and Evolution. 9 (3): 585–598. doi:10.1006/mpev.1998.0495. PMID 9668008.

[McKennaBell-6] McKenna, M.C.; Bell, S.K. (1997). Classification of mammals above the species level. New York: Columbia University Press. pp. 631 pp.

[Hughes-7] Hughes, K.; Elton, S.; O'Regan, J. (Jan 2008). "Theropithecus and 'Out of Africa' dispersal in the Plio-Pleistocene". Journal of Human Evolution. 54 (1): 43–77. doi:10.1016/j.jhevol.2007.06.004. ISSN 0047-2484. PMID 17868778.

[8] Maier, W. (1972). "The first complete skull of Simopithecus darti from Makapansgat, South Africa, and its systematic position". Journal of Human Evolution. 1 (4): 395–400. doi:10.1016/0047-2484(72)90116-9.

[Napier_1981-9] Napier PH. (1981). Catalogue of primates in the British museum (natural history) and elsewhere in the British Isles, part II: family Cercopithecidae, subfamily Cercopithecinae, London: British Museum (Natural History).

[Ankel-Simons_2007-10] ^ ^a ^b ^c ^d ^e ^f Ankel-Simons F. (2007). Primate Anatomy: an introduction, (3rd edition), San Diego: Elsevier Academic Press.

[Jolly_2007-11] Jolly CJ. (2007). "Baboons, mandrills, and mangabeys: afro-papionin socioecology in a phylogenetic perspective", In: Primates in perspective, Campbell CJ, Fuentes A, MacKinnon KC, Panger M, Bearder SK (eds), New York: Oxford U Pr. p 240-51.

[Dunbar_1977-12] ^ ^a ^b ^c ^d ^e ^f Dunbar RIM. (1977). "Feeding ecology of gelada baboons: a preliminary report", In: Primate ecology: studies of feeding and ranging behaviour in lemurs, monkeys and apes, Clutton-Brock TH (ed), London: Academic Pr. p 251-73.

[Iwamoto_1983-13] Iwamoto T, Dunbar RIM. (1983). "Thermoregulation, habitat quality and the behavioural ecology of gelada baboons", J Anim Ecol 52(2):357-66.

[14] Iwamoto, T. (1979). "Ecological and sociological studies of gelada. Feeding ecology". Contributions to Primatology. 16: 279–330. PMID 101341.

[Crook_1966-15] Crook JH. (1966). "Gelada baboon herd structure and movement: a comparative report", Symp Zool Soc Lond 18:237-58.

[Dunbar_1986-16] Dunbar RIM. (1986). "The social ecology of gelada baboons", In: Ecological aspects of social evolution: birds and mammals, Rubenstein DI, Wrangham RW, (eds), Primceton (NJ):Princeton U Pr. pp.332-51.

[Zinner_2004-17] Grüter CC, Zinner D. (2004). "Nested societies. Convergent adaptations of baboons and snub-nosed monkeys?" Prim Rep 70:1-98.

[Dunbar_1975-18] ^ ^a ^b ^c ^d ^e ^f Dunbar, R.; Dunbar, P. (1975). "Social dynamics of gelada baboons". Contributions to Primatology. 6: 1–157. PMID 808371.

[Dunbar_1980-19] Dunbar, R. I. M. (1980). "Determinants and evolutionary consequences of dominance among female gelada baboons". Behavioral Ecology and Sociobiology. 7 (4): 253–265. doi:10.1007/BF00300665.

[Ohsawa_1979-20] Ohsawa, H. (1979). "The local gelada population and environment of the Gich area". Contributions to Primatology. 16: 3–45. PMID 101342.

[male-female-21] Dunbar, R. I. M. (1983). "Structure of Gelada Baboon Reproductive Units III. The Males Relationship with his Females", Animal Behavior 31:565-575.

[Dunbar_1984-22] Dunbar, R. I. M. (1984). Reproductive decisions: an economic analysis of gelada baboon social strategies. Princeton, N.J: Princeton University Press. ISBN 978-0-691-08360-5.

[23] Roux, Aliza, Report, Department of Zoology and Entomology, Qwa Qwa campus, University of the Free State {{citation}}: Missing or empty |title= (help)

[24] "Wolves are better hunters when monkeys are around: An unexpected co-existence in the Ethiopian highlands". Science Daily. 22 June 2015.

[Bernstein_1975-25] Bernstein, I. S. (1975). "Activity patterns in a gelada monkey group". Folia Primatologica. 23 (1–2): 50–71. doi:10.1159/000155661. PMID 806510.

[behavior-26] ^ ^a ^b ^c ^d ^e ^f Mori, U. (1979). "Ecological and sociological studies of gelada baboons. Reproductive behavior". Contributions to Primatology. 16: 183–197. PMID 101336.

[Dunbar_2002-27] Dunbar, R. I. M.; Hannah-Stewart, L.; Dunbar, P. (2002). "Forage quality and the costs of lactation for female gelada baboons". Animal Behaviour. 64 (5): 801. doi:10.1006/anbe.2002.9972.

[28] Leutenegger, W. (1973). "Maternal-fetal weight relationships in primates". Folia Primatologica. 20 (4): 280–293. doi:10.1159/000155580. PMID 4208250.

[Barrett_1995-29] Barrett, L.; Dunbar, R. I. M.; Dunbar, P. (1995). "Mother-infant contact as contingent behaviour in gelada baboons". Animal Behaviour. 49 (3): 805. doi:10.1016/0003-3472(95)80211-8.

[30] "Gelada". nationalgeographic.com. National Geographic Society. 2011-05-10T06:34:00-0400. Retrieved 7 March 2012. {{cite web}}: Check date values in: |date= (help)

[Kawai_1979-31] Kawai, M. (1979). "Ecological and sociological studies of gelada baboons. Auditory communication and social relations". Contributions to Primatology. 16: 219–241. PMID 101338.

[32] Richman, Bruce (1976). "Some vocal distinctive features used by gelada monkeys". Journal of the Acoustical Society of America. 60 (3): 718–724. Bibcode:1976ASAJ...60..718R. doi:10.1121/1.381144. PMID 824335.

[unit2-33] Mori, U. (1979). "Ecological and sociological studies of gelada baboons. Individual relationships within a unit". Contributions to Primatology. 16: 93–124. PMID 101345.

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 | status = LC
 | status_system = IUCN3.1
-| status_ref = <ref name=iucn>{{Cite journal | author = Gippoliti, S. | author2 = Hunter, C. | last-author-amp = yes | title = ''Theropithecus gelada'' | journal = [[The IUCN Red List of Threatened Species]] | volume = 2008 | page = e.T21744A9316114 | publisher = [[IUCN]] | date = 2008 | url = http://www.iucnredlist.org/details/21744/0 | doi = 10.2305/IUCN.UK.2008.RLTS.T21744A9316114.en | access-date = 12 January 2018}}</ref>
+| status_ref = <ref name=iucn>{{Cite journal | author = Gippoliti, S. | author2 = Hunter, C. | last-author-amp = yes | title = ''Theropithecus gelada'' | journal = [[The IUCN Red List of Threatened Species]] | volume = 2008 | page = e.T21744A9316114 | date = 2008 | url = http://www.iucnredlist.org/details/21744/0 | doi = 10.2305/IUCN.UK.2008.RLTS.T21744A9316114.en | access-date = 12 January 2018}}</ref>
 | display_parents = 3
 | genus = Theropithecus
@@ Line 50: / Line 50: @@
 | last1 = Iwamoto | first1 = T.
 | title = Ecological and sociological studies of gelada. Feeding ecology
-| journal = Contributions to primatology
+| journal = Contributions to Primatology
 | volume = 16
 | pages = 279–330
@@ Line 68: / Line 68: @@
 | last1 = Ohsawa | first1 = H.
 | title = The local gelada population and environment of the Gich area
-| journal = Contributions to primatology
+| journal = Contributions to Primatology
 | volume = 16
 | pages = 3–45
@@ Line 77: / Line 77: @@
 | last2 = Dunbar | first2 = P.
 | title = Social dynamics of gelada baboons
-| journal = Contributions to primatology
+| journal = Contributions to Primatology
 | volume = 6
 | pages = 1–157
@@ Line 87: / Line 87: @@
 Males can remain in a reproductive unit for four to five years.<ref name= "Dunbar 1986"/> While geladas have traditionally been considered to have a male-transfer society, many males appear to be likely to return and breed in their natal bands. Nevertheless, gelada males leave their natal units and try to take over a unit of their own. A male can take over a reproductive unit either through direct aggression and fighting or by joining one as a subordinate and taking some females with him to create a new unit.<ref name= "Dunbar 1986"/> When more than one male is in a unit, only one of them can mate with the females.<ref name= "Dunbar 1975"/><ref name= "Ohsawa 1979"/> The females in the group together can have power over the dominant male. When a new male tries to take over a unit and overthrow the resident male, the females can choose to support or oppose him. The male maintains his relationship with the females by grooming them rather than forcing his dominance, in contrast to the society of the hamadryas baboon. Females accept a male into the unit by presenting themselves to him. Not all the females may interact with the male. Usually, one may serve as his main partner.<ref name= "male-female">Dunbar, R. I. M. (1983). "Structure of Gelada Baboon Reproductive Units III. The Males Relationship with his Females", ''Animal Behavior'' 31:565-575.</ref> The male may sometimes be monopolized by this female.<ref name= "male-female"/> The male may try to interact with the other females, but they are usually unresponsive.<ref name= "male-female"/>
-Most all-male units consist of several subadults and one young adult, led by one male. A member of an all-male unit may spend two to four years in the group before attempting to join a reproductive unit. All-male groups are generally aggressive towards both reproductive units and other all-male units.<ref name= "Dunbar 1975"/> As in reproductive units, aggression within all-male units is rare. As bands, reproductive units exist in a common home range.<ref name= "Dunbar 1984">{{cite book |last= Dunbar |first=R. I. M. |title= Reproductive decisions: an economic analysis of gelada baboon social strategies |publisher= Princeton University Press |location=Princeton, N.J |year=1984 |pages= |isbn=0-691-08360-6 |oclc= |doi= |accessdate=}}</ref> Within the band, members are closely related and between the units there is no social hierarchy. Bands usually break apart every eight to nine years as a new band forms in a new home range.
+Most all-male units consist of several subadults and one young adult, led by one male. A member of an all-male unit may spend two to four years in the group before attempting to join a reproductive unit. All-male groups are generally aggressive towards both reproductive units and other all-male units.<ref name= "Dunbar 1975"/> As in reproductive units, aggression within all-male units is rare. As bands, reproductive units exist in a common home range.<ref name= "Dunbar 1984">{{cite book |last= Dunbar |first=R. I. M. |title= Reproductive decisions: an economic analysis of gelada baboon social strategies |publisher= Princeton University Press |location=Princeton, N.J |year=1984 |pages= |isbn=978-0-691-08360-5 |oclc= |doi= }}</ref> Within the band, members are closely related and between the units there is no social hierarchy. Bands usually break apart every eight to nine years as a new band forms in a new home range.
 Researchers from the [[University of the Free State]] (UFS) in [[South Africa]], while observing gelada during field studies, discovered that the monkeys were capable of 'cheating' on their partners and covering up their 'infidelity'. A non-dominant male would mate surreptitiously with a female, suppressing their normal mating cries so as not to be overheard. If discovered, the dominant male would attack the miscreants in a clear form of punishment. It is the first time that evidence of the knowledge of cheating and fear of discovery has been recorded among animals in the wild. Dr. Aliza le Roux of the university's Department of Zoology and Entomology believes that dishonesty and punishment are not uniquely human traits, and that the observed evidence of this behaviour among gelada monkeys suggests that the roots of the human system of deceit, crime and punishment lie very deep indeed.<ref>{{cite|first=Aliza |last=le Roux| work= Report |publisher= Department of Zoology and Entomology, Qwa Qwa campus, [[University of the Free State]]}}</ref>
-Mixed-species association was observed between solitary [[Ethiopian wolf|Ethiopian wolves]] and geladas. According to the study's findings, gelada monkeys would not typically move on encountering Ethiopian wolves, even when they were in the middle of the herd—68 percent of encounters resulted in no movement and only 11 percent resulted in a movement of greater than {{cvt|10|m}}. In stark contrast, the geladas always fled great distances to the cliffs for safety whenever they encountered aggressive domestic dogs.<ref>{{cite journal |title= Wolves are better hunters when monkeys are around: An unexpected co-existence in the Ethiopian highlands | work=Science Daily | date=22 June 2015| url= https://www.sciencedaily.com/releases/2015/06/150622085224.htm}}</ref>
+Mixed-species association was observed between solitary [[Ethiopian wolf|Ethiopian wolves]] and geladas. According to the study's findings, gelada monkeys would not typically move on encountering Ethiopian wolves, even when they were in the middle of the herd—68 percent of encounters resulted in no movement and only 11 percent resulted in a movement of greater than {{cvt|10|m}}. In stark contrast, the geladas always fled great distances to the cliffs for safety whenever they encountered aggressive domestic dogs.<ref>{{cite journal |title= Wolves are better hunters when monkeys are around: An unexpected co-existence in the Ethiopian highlands | journal=Science Daily | date=22 June 2015| url= https://www.sciencedaily.com/releases/2015/06/150622085224.htm}}</ref>
 ===Reproduction and parenting===
@@ Line 108: / Line 108: @@
 | last1 = Mori | first1 = U.
 | title = Ecological and sociological studies of gelada baboons. Reproductive behavior
-| journal = Contributions to primatology
+| journal = Contributions to Primatology
 | volume = 16
 | pages = 183–197
@@ Line 125: / Line 125: @@
 | year = 1973
 | pmid = 4208250
-}}</ref> Females that have just given birth stay on the periphery of the reproductive unit. Other adult females may take an interest in the infants and even kidnap them.<ref name= "behavior"/> An infant is carried on its mother’s belly for the first five weeks, and thereafter on her back.<ref name= "behavior"/><ref name= "Barrett 1995">{{Cite journal | last1 = Barrett | first1 = L. | last2 = Dunbar | first2 = R. I. M. | authorlink2 = Robin Dunbar| last3 = Dunbar | first3 = P. | doi = 10.1016/0003-3472(95)80211-8 | title = Mother-infant contact as contingent behaviour in gelada baboons | journal = Animal Behaviour | volume = 49 | issue = 3 | pages = 805 | year = 1995 | pmid =  | pmc = }}</ref> Infants can move independently at around five months old. A subordinate male in a reproductive unit may help care for an infant when it is six months old.<ref name= "behavior"/> When herds form, juveniles and infants may gather into play groups of around ten individuals. When males reach puberty, they gather into unstable groups independent of the reproductive units. Females sexually mature at around three years, but do not give birth for another year.<ref name= "Dunbar 1975"/><ref name= "Dunbar 1984"/> Males reach puberty at about four or five years, but they are usually unable to reproduce because of social constraints and wait until they are about eight to ten years old.<ref name= "Jolly 2007"/> Average life span in the wild is 19 years.<ref>{{cite web|title= Gelada| website= nationalgeographic.com |url= http://animals.nationalgeographic.com/animals/mammals/gelada/ |publisher=[[National Geographic Society]]|accessdate=7 March 2012}}</ref>
+}}</ref> Females that have just given birth stay on the periphery of the reproductive unit. Other adult females may take an interest in the infants and even kidnap them.<ref name= "behavior"/> An infant is carried on its mother’s belly for the first five weeks, and thereafter on her back.<ref name= "behavior"/><ref name= "Barrett 1995">{{Cite journal | last1 = Barrett | first1 = L. | last2 = Dunbar | first2 = R. I. M. | authorlink2 = Robin Dunbar| last3 = Dunbar | first3 = P. | doi = 10.1016/0003-3472(95)80211-8 | title = Mother-infant contact as contingent behaviour in gelada baboons | journal = Animal Behaviour | volume = 49 | issue = 3 | pages = 805 | year = 1995 | pmid =  | pmc = }}</ref> Infants can move independently at around five months old. A subordinate male in a reproductive unit may help care for an infant when it is six months old.<ref name= "behavior"/> When herds form, juveniles and infants may gather into play groups of around ten individuals. When males reach puberty, they gather into unstable groups independent of the reproductive units. Females sexually mature at around three years, but do not give birth for another year.<ref name= "Dunbar 1975"/><ref name= "Dunbar 1984"/> Males reach puberty at about four or five years, but they are usually unable to reproduce because of social constraints and wait until they are about eight to ten years old.<ref name= "Jolly 2007"/> Average life span in the wild is 19 years.<ref>{{cite web|title= Gelada| website= nationalgeographic.com |url= http://animals.nationalgeographic.com/animals/mammals/gelada/ |publisher=[[National Geographic Society]]|accessdate=7 March 2012| date= 2011-05-10T06:34:00-0400 }}</ref>
 ===Communication===
@@ Line 131: / Line 131: @@
 | last1 = Kawai | first1 = M.
 | title = Ecological and sociological studies of gelada baboons. Auditory communication and social relations
-| journal = Contributions to primatology
+| journal = Contributions to Primatology
 | volume = 16
 | pages = 219–241
@@ Line 139: / Line 139: @@
 | last1 = Mori | first1 = U.
 | title = Ecological and sociological studies of gelada baboons. Individual relationships within a unit
-| journal = Contributions to primatology
+| journal = Contributions to Primatology
 | volume = 16
 | pages = 93–124

Taxon identifiers
Theropithecus gelada	Wikidata: Q194080 Wikispecies: Theropithecus gelada ADW: Theropithecus_gelada ARKive: theropithecus-gelada BOLD: 12481 CoL: 7C8JR ECOS: 4110 EoL: 311178 GBIF: 2436553 iNaturalist: 43530 IRMNG: 11055292 ITIS: 573034 IUCN: 21744 MDD: 1000635 MSW: 12100593 NCBI: 9565 Observation.org: 88041 Open Tree of Life: 741053 Paleobiology Database: 233450 Species+: 3493
Macacus gelada	Wikidata: Q41165800 GBIF: 9017781 ZooBank: 8C9CFE1C-0352-4F52-BDBF-B0CD0F4BC1B1