Mantled guereza

Mantled guereza[1]
with baby at Münster Zoo
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Clade: Synapsida
Class: Mammalia
Order: Primates
Suborder: Haplorhini
Family: Cercopithecidae
Genus: Colobus
Species: C. guereza
Binomial name
Colobus guereza
Rüppell, 1835
Mantled guereza range

The mantled guereza (Colobus guereza), also known simply as the guereza, the eastern black-and-white colobus, or the Abyssinian black-and-white colobus, is a black-and-white colobus, a type of Old World monkey. It is native to much of west central and east Africa, including Cameroon, Equatorial Guinea, Nigeria, Ethiopia, Kenya, Tanzania, Uganda and Chad. The species consists of several subspecies that differ in appearance. It has a distinctive appearance, which is alluded to in its name; the long white fringes of hair the run along each side of its black trunk are known as a mantle. Its face is framed with white hair and it has a large white tail tuft.

The mantled guereza is diurnal and arboreal, found in both deciduous and evergreen forests. It is an adaptable species that can cope with habitat disturbance and prefers secondary forest close to rivers or lakes. Although previously thought only to eat leaves, it also eats seeds, fruits, and arthropods. It is able to digest plant material with a high fibre content with its specialised stomach and may only eat from a few plant species at a time. It is preyed on by birds of prey and some mammals, such as the common chimpanzee and the leopard.

The mantled guereza lives in social groups of three to fifteen individuals. These groups normally include a dominant male, several females, and the offspring of the females. It has a polygynous mating system and copulation is initiated with vocal communication. After a gestation period of just over five months, infants are born with pink skin and white fur, which darkens to the adult coloration by three to four months. The mantled guereza is well known for its dawn chorus, the males' "roar" is a method of long distance communication that reinforces territorial boundaries. It also makes other vocalization and uses body postures, movements, and facial expressions to communicate.

The mantled guereza is listed as Least Concern by the International Union for Conservation of Nature (IUCN) because it is widespread - although it is locally threatened in some areas, the decline is not great enough to list it in a higher category of threat. However, one subspecies found in Kenya is listed as Endangered. It can survive well in degraded forests and in some areas it is more common in logged areas than unlogged ones. The mantled guereza is also threatened by hunting for bushmeat and for its skin.

Etymology

The mantled guereza has many alternative common names including the guereza, the eastern black-and-white colobus, the magistrate colobus,[2] or the Abyssinian black-and-white colobus.[3] The name "mantled" refers to its mantle, the long silky white fringes of hair that run along its body and "guereza" is the native name of the monkey in Ethiopia.[4] The scientific name Colobus derives from Greek kolobus meaning "mutilated" which refers to its lack of thumbs.[5]

Discovery and taxonomic classification

The mantled guereza was discovered by Eduard Rüppell, a German naturalist and explorer, during his trip to Abyssinia between 1830 and 1834.[6] He wrote about the species in Neue Wirbelthiere con Abyssinien, Saengthiere in 1835.[7] It was first seen in Europe in 1890 in Berlin Zoological Garden when three individuals were purchased from a dealer from Massawa, Eritrea.[8]

The mantled guereza is in the Colobinae subfamily, also known as the leaf-eating monkeys, a group of Old World monkeys from Asia and Africa. This subfamily is split into three groups, the colobus monkeys of Africa, of which the mantled guereza is a part, the langurs, or leaf monkeys, of Asia, and an "odd-nosed" group. The African colobus monkeys are divided again by distinctions in color, behavior, and ecology. The three genera are the black-and-white colobi, the red colobi, and the olive colobi. There are three black-and-white colobi: the mantled guereza, Colobus guereza, the king colobus, C. polykomos, and the Angola colobus, C. angolensis.[9] Groves lists seven subspecies of mantled guereza in Mammal Species of the World (MSW) (2005).[1] The validity of the Djaffa Mountain guereza, C. g. gallarum, is uncertain, although not listed by Groves in MSW, it is recognised in his 2007 Colobinae review paper,[10] and by International Union for Conservation of Nature (IUCN) assessors Gippolliti and Butynski in 2008.[2]

The morphological difference between subspecies is most pronounced between the southeastern Kilimanjaro guereza, C. g. caudatus, and the northwestern western guereza, C. g. occidentalis. The intermediate subspecies show a gradual change between the two.[10]

Physical description

Mantled guereza portrait showing its facial features

The mantled guereza has a distinctive pelage, it is mostly black, with long white fringes of silky hair—known as a mantle or ornamentation—along the sides of its body and tail. The bands that make up the mantle start at the shoulders and extend along the back until they connect at the lower torso. The tail is long and ends in a white tuft which varies in how much it covers the tail. These features vary in color among subspecies, for example the tail of C. g. guereza is gray until the white tail tuft which covers half of its length, while the tail tuft of C. g. caudatus makes up 80% of the tail. The mantle color ranges from white to cream or yellow. Its face is framed by white hair and it has bushy cheek hairs. The thigh has a white stripe.[11]

Infants are born with pink skin and white hair. The hair and skin darken as they age and by three to four months they attain adult coloration. Male usually gain their coloration before females.[12] The male typically weighs 9.3 and 13.5 kilograms (21 and 30 lb) and the female weighs between 7.8 and 9.2 kilograms (17 and 20 lb). The head and body length averages 61.5 centimetres (24.2 in) for males and 57.6 centimetres (22.7 in) for females. Like most colobi, the mantled guereza has a small thumb that is vestigial.[13][14] There is dentition sexual dimorphism among the subspecies. In some, the males have larger teeth than females, in others the reverse is true, and some have no significant difference.[15]

Distribution and habitat

Mantled guerezas are arboreal and prefer secondary forests.

The mantled guereza is distributed throughout Equatorial Africa. It ranges from Nigeria and Cameroon in the west to Ethiopia, Kenya, Uganda, and northern Tanzania in the east.[11][13] The mantled guereza lives in both deciduous and evergreen forests. It mainly inhabits forest and savannah woodlands and often extend into highland and montane forests.[14] It can be found in other forest habitats, both primary and secondary, such as riparian (near fresh or brackish water), gallery, and upland forests. It is particularly common in forests close to rivers and lakes and at high elevations.[16] It can be found in elevations as high as 3,300 metres (10,800 ft).[17] This species prefers secondary forests and selects them over old-growth forests if given the choice.[18] It is likely that the mantled guereza prefers these forests due to the increased number of food trees and the weaker chemical defenses of the species within.[18] The mantled guereza is sometimes found in swamps[19] as well as human-made habitats such as Eucalyptus plantations, which may be frequented when the monkey has nutritional deficiencies.[17]

Ecology

The mantled guereza is primarily arboreal, but does sometimes descend on the ground to forage and travel, perhaps more so than most other colobines. It is diurnal and rests for up to half the day. Foraging or travelling are the next most common activity. Sometime after dawn, mantled guereza groups leave their sleeping trees and will return to them at dusk. During the day, the mantled guereza has long rest periods in between periods of moving and feeding.[20][21][22] Other activities, including grooming, greeting, playing and being vigilant, are performed to a lesser extent.[21][22]

The diet of the mantled guereza is predominantly leaves, often of only a few tree species.

Despite its reputation as an exclusive leaf-eater, the mantled guereza is not an obligate folivore.[14] While it mainly eats leaves and fruit, its diet is quite variable. It may eat bark, wood, seeds, flowers, petioles, lianas, aquatic-plants, arthropods, soil, and even concrete from buildings.[23] The amount of each food item in its diet varies by area and time of year. Nutritional factors like protein, tannins, and sodium levels in leaves influence its food choices. It may even intermittently travel longer distances to access plants with higher levels of nutrition.[24] Leaves usually make up over half of its diet, although fruits are occasionally eaten more depending on the season.[14][17] When foraging for leaves, the mantled guereza prefers young ones over old.[18] With fleshy fruits, the mantled guereza prefers to eat them unripe, which may serve to reduce competition with primates that eat ripe fruits.[23] It consumes a number of plant species but only some make up most of its diet at a specific site.[19][23]

Like all colobi, the mantled guereza is able to digest leaves and other plant fibers with a large, multi-chambered stomach that contains bacteria in certain areas.[25] Like most colobines, it prefers foods with high fiber content that can be easily extracted with its specialized stomach.[24] The mantled guereza is mostly preyed on by the crowned hawk-eagle,[26] but it is also eaten by other birds of prey such as Verreaux's eagle.[17] The common chimpanzee is known to hunt the guereza.[27] The leopard is another possible predator.[28]

Behaviour

Social structure

Social grooming or allogrooming mainly occurs between females and is an important social interaction in mantled guereza groups.

The mantled guereza lives in stable social groups usually containing three to fifteen members.[21] The groups usually contain one male, several females and juveniles. In some populations, groups containing several males are common.[29] In multi-male groups, males tend to be aggressive with one another with one being dominant. Some males may be expelled from these groups.[20] Multi-male groups may contain father-son pairs or unrelated males.[30] Males that are not part of groups either live solitarily or with other outside males in bachelor groups. The females keep the groups cohesive and they are matrilineally related. They rarely disperse from their natal groups, except possibly when they break apart.[20] Males on the other hand, usually leave when they become subadults or adults. They may start out being solitary and or in bachelor groups. They gain entry into a social group either by being on the periphery or displacing a group male.[17]

Because of its low quality diet and the dispersed distribution of its food, the mantled guereza has a resident-egalitarian social structure. Female guerezas living in a group often have an egalitarian dominance style with no formalized rank relations. Relationships are relaxed and friendly with rare signals of dominance or subordinance. Physical aggression within the group is usually not harmful and rarely escalates into a conflict. Allogrooming is an important part of mantled guereza interactions and mostly occurs between females.[31] The adult males rarely groom in the groups. While not strictly territorial, mantled guereza groups can be aggressive towards each other.[21] In some populations, groups may defend core areas (which exist as a small part of the home range), resources, and mates. During intergroup encounters, males can engage in direct or indirect mate defense, like defending a female's resources.[32] It is the males that participate in agonistic inter-group encounters but female may do so as well.[21] Aggressive encounters between groups usually involve chases, displays and vocalizations rather than physical contact.[21]

Reproduction and parenting

Infants are born with white fur and are always carried during the first months of their life.

The mantled guereza has a polygynous harem-based mating system.[14][20] Mating solicitations are made by both males and females, half of the time for each.[33] To solicit mating, the mantled guereza will walk near its potential partner and make low-intensity mouth clicks or tough-smacks.[34] During copulation, the males hold on the female's ankles and body.[35] Most matings take place between individuals of the same group but copulations outside of the group have been recorded.[35] In multi-male groups, more than one male may mate with the females.[21] The gestation period lasts 158 days with a 16-22 month interbirth interval.[17] The newborn guereza relies on its mother for support and must cling to her. As they grow older, infants can move on their own but keep returning to their mothers.[36] The infants take up most of the attention in the groups. The other females in a group may handle an infant although the latter are only comfortable with their mothers.[37] The males normally don't pay much attention to infants until they are four to five weeks old.[36] Infants can eat solid food at about eight to nine weeks and by fifty weeks they are fully weaned and no longer need to hold on to their mothers.[37]

Communication

The most notable vocalization of the mantled guereza is the "roar", which is made mainly at night or dawn by males. The sound of a roar can be carried for up to a mile. It is normally the dominant male who roars when there are multiple males in the group. Roars are used for long distance communication and can regulate inter-group spacing without direct, physical contact while foraging.[38] When one male starts roaring, neighboring males will start to roar as well.[35] Often, the mantled guereza will respond to calls regardless of "caller identity," focusing more on the collective vocal displays and not the familiarity of the caller.[38] There is variation in the roars of males which could signal the status of their group and fighting ability.[35] With a roar, a male can advertise his body size; both actual and exaggerated.[39] Other vocalizations are made as well. Males may snort, possibly as an alarm call. "Purrs" are made before group movements. Females and infants may "caw" when under mild distress. When in more serious distress, like if an infant is in danger, females and sub-adults will squeak or scream. "Tongue-clicking" is made during mild aggression.[40] In addition to vocalizations, the mantled guereza communicates with several different body postures and movements, displaying of fringe fur, facial expressions, and touches.[37]

Conservation status

The mantled guereza is hunted for its skin and meat.

Because it can live in both dry and gallery forests and move on the ground, the mantled guereza is less threatened than many other colobine species.[14] The IUCN lists it as Least Concern because "although locally threatened in parts of its range, this widespread species is not thought to be declining fast enough to place it in a higher category of threat."[2] However, some of the subspecies are classified under different categories. The Mt Uaraguess guereza, C. g. percivali, is listed as Endangered due to its small range and its risk from hunting,[41] while the Dodinga Hills guereza, C. g. dodingae, the Djaffa Mountains guereza, C. g. gallarum, and the Mau Forest guereza, C. g. matschiei are all listed as Data Deficient.[42][43][44]

The Dodinga Hills guereza hasn't been recorded since the 1960s.[42] Unlike most other primate species, the mantled guereza can survive habitat degradation and can even thrive in degraded forests.[45] Sometimes, logging may increase the number of preferred food trees for the mantled guereza and it is more abundant in logged areas than unlogged ones.[14] However, complete forest clearance causes dramatic declines in numbers. In Uganda complete forest clearings caused a decline of 50% over eight years.[46] The mantled guereza is also threatened by hunting for meat and its skin. Mantled guereza meat sells as bushmeat for $4–9 US.[47] The skins have been sold for fashion or in the tourist trade.[48]

References

  1. 1 2 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. 168. OCLC 62265494. ISBN 0-801-88221-4.
  2. 1 2 3 4 Kingdon, J.; Struhsaker, T.; Oates, J. F.; Hart, J.; Groves, C. P. (2008). Colobus guereza. In: IUCN 2008. IUCN Red List of Threatened Species. Retrieved 2 April 2013.
  3. ↑ Wolfheim, J. H. (1983). Primates Of The World: Distribution, Abundance And Conservation. Routledge. ISBN 3-7186-0190-7.
  4. ↑ "Guereza". Merriam-Webster dictionary. Merriam-Webster. Retrieved 2 April 2013.
  5. ↑ Grzimek, B. (1972). Grzimek's Animal Life Encyclopedia: Mammals I-IV. Volume 10. Van Nostrand Reinhold Co. p. 464.
  6. ↑ Beolens, B.; Watkins, M.; Grayson, M. (2009). The Eponym Dictionary of Mammals. JHU Press. p. 353. ISBN 978-0-8018-9533-3.
  7. ↑ A Catalogue of the Mammalia in the Museum of the Hon. East-India Company (Google eBook). Allen. 1851. p. 16.
  8. ↑ "Popular Science" (Google eBook) 41 (22). 1892: 393. ISSN 0161-7370.
  9. ↑ Fleagle, J. G. (1998). Primate Adaptation and Evolution (2nd ed.). Academic Press. pp. 207–209. ISBN 978-0-12-260341-9.
  10. 1 2 Groves, C. (2007). "The taxonomic diversity of the Colobinae of Africa" (PDF). Journal of Anthropological Sciences 85: 7–34.
  11. 1 2 Groves, C. (2001). Primate Taxonomy. Washington DC: Smithsonian Institution Press.
  12. ↑ Ackerman, D. E. (1991). "A study of the colobus monkey (Colobus guereza kikuyuensis)". Animal Keeper's Forum 18 (4): 164–171.
  13. 1 2 Napier, P. H. (1985). Catalogue of Primates in the British Museum (Natural History) and Elsewhere in the British Isles, part III: Family Cercopithecidae, Subfamily Colobinae. London: British Museum (Natural History).
  14. 1 2 3 4 5 6 7 Oates, J. F. (1994). "The natural history of African colobines". In Davies, A. G.; Oates, J. F. Colobine Monkeys: Their Ecology, Behaviour and Evolution. Cambridge, UK: Cambridge University Press. pp. 75–128.
  15. ↑ Hayes, V. J.; Freedman, L.; Oxnard, C. E. (1995). "The differential expression of dental sexual dimorphism in subspecies of Colobus guereza". International Journal of Primatology 16 (6): 971–996. doi:10.1007/bf02696112.
  16. ↑ Dunbar, R. I. M. (1987). "Habitat quality, population dynamics, and group composition in colobus monkeys (Colobus guereza)". International Journal of Primatology 8 (4): 299–329. doi:10.1007/bf02737386.
  17. 1 2 3 4 5 6 Dunbar, R. I. M.; Dunbar, E. P. (1974). "Ecology and population dynamics of Colobus guereza in Ethiopia". Folia Primatologica 21: 188–208. doi:10.1159/000155600.
  18. 1 2 3 Lwanga, J. S. (2006). "Spatial distribution of primates in a mosaic of colonizing and old growth forest at Ngogo, Kibale National Park, Uganda" (PDF). Primates 47 (3): 230–238. doi:10.1007/s10329-005-0173-5. PMID 16479411.
  19. 1 2 Oates, J. F. (1978). "Water-plant and soil consumption by guereza monkeys (Colobus guereza): a relationship with minerals and toxins in the diet?". Biotropica 10 (4): 241–253. doi:10.2307/2387676.
  20. 1 2 3 4 Bocian, C. M. (1997). Niche separation of black-and-white colobus monkeys (Colobus angolensis and C. guereza) in the Ituri Forest (Ph.D.). City University of New York.
  21. 1 2 3 4 5 6 7 von Hippel, F. A. (1996). "Interactions between overlapping multimale groups of black and white colobus monkeys (Colobus guereza) in the Kakamega Forest, Kenya". American Journal of Primatology 38 (3): 193–209. doi:10.1002/(SICI)1098-2345(1996)38:3<193::AID-AJP1>3.0.CO;2-U.
  22. 1 2 Oates, J. F. (1977a). "The guereza and its food". In Clutton-Brock, T. H. Primate Ecology: Studies of Feeding and Ranging Behaviour in Lemurs, Monkeys and Apes. London: Academic Press. pp. 275–321.
  23. 1 2 3 Harris, T. R.; Chapman, C. A. (2007). "Variation in diet and ranging of black and white colobus monkeys in Kibale National Park, Uganda" (PDF). Primates 48 (3): 208–221. doi:10.1007/s10329-006-0036-8. PMID 17429575.
  24. 1 2 Fashing, P. J.; Dierenfeld, E. S.; Mowry, C. B. (2007). "Influence of plant and soil chemistry on food selection, ranging patterns, and biomass of Colobus guereza in Kakamega Forest, Kenya" (PDF). International Journal of Primatology 28 (3): 673. doi:10.1007/s10764-006-9096-2.
  25. ↑ Oates, J. F.; Davies, A. G.; Delson, E (1994). "The diversity of living colobines". In Davies, A. G.; Oates, J. F. Colobine Monkeys: Their Ecology, Behaviour and Evolution. Cambridge, UK: Cambridge University Press. pp. 45–73.
  26. ↑ Struhsaker, T. T.; Leakey, M. (1990). "Prey selectivity by crowned hawk-eagles on monkeys in the Kibale Forest, Uganda". Behavioral Ecology and Sociobiology 26 (6): 435–443. doi:10.1007/bf00170902.
  27. ↑ Ihobe, H. (2001). "Hunting attempt by chimpanzees on Abyssinian colobus at the Kalinzu Forest, Uganda". Pan African News 8 (2): 31–32.
  28. ↑ Schel, A. M.; Tranquilli, S.; Zuberbühler, K. (2009). "The alarm call system of two species of black-and-white colobus monkeys (Colobus polykomos and Colobus guereza)". Journal of Comparative Psychology 123 (2): 136–150. doi:10.1037/a0014280. PMID 19450021.
  29. ↑ Dunbar, R. I. M. (1976). "Contrasts in social structure among black-and-white colobus monkey groups". Animal Behaviour 24 (1): 84–92. doi:10.1016/s0003-3472(76)80102-9.
  30. ↑ Harris, T. R.; Caillaud, D.; Chapman, C. A.; Vigilant, L. (2009). "Neither genetic nor observational data alone are sufficient for understanding sex-biased dispersal in a social-group-living species". Molecular Ecology 18: 1777–1790. doi:10.1111/j.1365-294x.2009.04139.x.
  31. ↑ Grunau, T.; Kuester, J. (2001). "Dominance style in female guerezas (Colobus guereza Rüppell 1835)". Primates 42 (4): 301. doi:10.1007/BF02629621.
  32. ↑ Fashing, P. (2001). "Male and female strategies during intergroup encounters in guerezas (Colobus guereza): Evidence for resource defense mediated through males and a comparison with other primates" (PDF). Behavioral Ecology and Sociobiology 50 (3): 219–230. doi:10.1007/s002650100358.
  33. ↑ Harris, T. R.; Monfort, S. L. (2006). "Mating behavior and endocrine profiles of wild black and white colobus monkeys (Colobus guereza): Toward an understanding of their life history and mating system". American Journal of Primatology 68 (4): 383–396. doi:10.1002/ajp.20232. PMID 16534807.
  34. ↑ Grimes, K. H. (2000). Guereza dietary and behavioural patterns at the Entebbe Botanical Gardens (M.A.). University of Calgary.
  35. 1 2 3 4 Harris, T. R. (2005). Roaring, intergroup aggression, and feeding competition in black and white colobus monkeys (Colobus guereza) at Kanyawara, Kibale National Park, Uganda (Ph.D.). Yale University.
  36. 1 2 Horwich, R. H.; Manski, D. (1975). "Maternal care and infant transfer in two species of Colobus monkeys" (PDF). Primates 16: 49. doi:10.1007/BF02381799.
  37. 1 2 3 Oates, J. F. (1977). "The social life of a black-and-white colobus monkey, Colobus guereza". Zeitschrift für Tierpsychologie 45: 1–0. doi:10.1111/j.1439-0310.1977.tb01007.x.
  38. 1 2 Schel, A. M.; Zuberbühler, K. (2011). "Dawn chorusing in guereza colobus monkeys". Behavioral Ecology and Sociobiology 66 (3): 361. doi:10.1007/s00265-011-1282-3.
  39. ↑ Harris, T. R.; Fitch, W. T.; Goldstein, L. M.; Fashing, P. J. (2006). "Black and white colobus monkey (Colobus guereza) roars as a source of both honest and exaggerated information about body mass" (PDF). Ethology 112 (9): 911. doi:10.1111/j.1439-0310.2006.01247.x.
  40. ↑ Marler, P. (1972). "Vocalizations of East African monkeys II: black and white colobus". Behaviour 42 (3): 175. doi:10.1163/156853972X00266.
  41. ↑ Struhsaker, T.; Oates, J. F.; Hart, J.; Butynski, T. M.; Groves, C. P. (2008). Colobus guereza ssp. percivali. In: IUCN 2008. IUCN Red List of Threatened Species. Retrieved 2 April 2013.
  42. 1 2 Struhsaker, T.; Oates, J. F.; Hart, J.; Butynski, T. M.; Groves, C. P. (2008). Colobus guereza ssp. dodingae. In: IUCN 2008. IUCN Red List of Threatened Species. Retrieved 4 April 2013.
  43. ↑ Gippolliti, S; Butynski, T. M. (2008). Colobus guereza ssp. gallarum. In: IUCN 2008. IUCN Red List of Threatened Species. Retrieved 4 April 2013.
  44. ↑ Struhsaker, T.; Oates, J. F.; Hart, J.; Butynski, T. M.; Groves, C. P. (2008). Colobus guereza ssp. matschiei. In: IUCN 2008. IUCN Red List of Threatened Species. Retrieved 4 April 2013.
  45. ↑ Chapman, C. A.; Balcomb, S. R.; Gillespie, T. R.; Skorupa, J. P.; Struhsaker, T. T. (2000). "Long-term effects of logging on African primate communities: A 28-year comparison from Kibale National Park, Uganda" (PDF). Conservation Biology 14: 207. doi:10.1046/j.1523-1739.2000.98592.x.
  46. ↑ Chapman, C. A.; Naughton-Treves, L.; Lawes, M. J.; Wasserman, M. D.; Gillespie, T. R. (2007). "Population declines of Colobus in western Uganda and conservation value of forest fragments" (PDF). International Journal of Primatology 28 (3): 513. doi:10.1007/s10764-007-9142-8.
  47. ↑ Eves, H. E.; Ruggiero, R. G. (2000). "Socioeconomics and the sustainability of hunting in the forests of northern Congo (Brazzaville)". In Robinson, J. G.; Bennett, E. L. Hunting for Sustainability in Tropical Forests. New York: Columbia University Press. pp. 427–454. ISBN 978-0-231-50492-8.
  48. ↑ Oates, J. F. (1977c). "The guereza and man". In Rainier III, Prince of Monaco; Bourne, G. H. Primate Conservation. New York: Academic Press. pp. 419–467. ISBN 978-0-12-576150-5.

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