Cercopithecidae Author Russell A. Mittermeier Author Anthony B. Rylands Author Don E. Wilson text 2013 2013-03-31 Lynx Edicions Barcelona Handbook of the Mammals of the World – Volume 3 Primates 550 755 book chapter 100956 10.5281/zenodo.6867065 3d520847-5163-4b5c-87bf-2cdceb781098 978-84-96553-89-7 6867065 82. Angolan Colobus Colobus angolensis French: Colobe d'Angola / German: Angola-Stummelaffe / Spanish: Colobo de Angola Other common names: Angola Colobus , Angolan Black-and-white Colobus; Adolf Friedrich’s Angolan/Ruwenzori Black-and-white Colobus (ruwenzorii) , Cordier’'s Angolan Colobus (cordieri) , Peters’'s Angolan/Tanzanian Black-andwhite Colobus (palliatus) , Powell-Cotton’s Angolan Colobus (cottoni) , Prigogine’s Angolan/Prigogine’s Black-and-white Colobus (prigoginel) , Sclater's Angolan Colobus (angolensis) , Sharpe’s/Southern Highlands Angolan Colobus (sharpe)) Taxonomy. Colobus angolensis Sclater, 1860 , Angola, 300 miles (483 km) inland from Bembe. In 2001, C. P. Groves recognized all of the named subspecies listed here except for sharper, which he believed to be a junior synonym of palliatus. C. sharpei was considered a valid subspecies of C. angolensis by U. Rahm in 1970, J. Kingdon in his 1971 review of East African primates, and P. Napier in 1985. In 1981, W. Rodgers provided information on the distributions of palliatus and sharpei that showed they were not isolated as they were thought to be and suggested they were not distinct taxa. Nevertheless, a molecular genetic study of palliatus across its distribution by M. McDonald and H. Hamilton in 2010 demonstrated distinct haplotypes in Kenyan and Tanzanian populations, which, combined with pelage differences highlighted by Rahm, Napier, and Kingdon, supported the validity of sharpei in central and southern Tanzania as a subspecies distinct from palliatus in Kenya. The form prigogine : could be synonymous with cordieri, but it is recognized as a subspecies of C. angolensis here. An unnamed subspecies of C. angolensis , the “Nkungwe’s Angolan Colobus ” or “Mahale Mountains Angola Colobus ” was listed by Groves in 2001 and P. Grubb and colleagues in their taxonomic review in 2003. It is an isolated population reported by T. Nishida and colleagues in 1981 from Mount Nkwunge and ridges of the northern Mahale Mountains as far as Mount Pasagula to the north and Mount Kahoko and Mount Sibindi to the south. It wasillustrated and described in a field guide of primates of the Mahale Mountains by T. Butynski and Y. de Jong in 2009, but it remains without a formal scientific trinomial. Seven subspecies recognized. Subspecies and Distribution. C.a.angolensisSclater,1860—DRCongo(SofthegreatbendoftheCongoRiver),NEAngola,andNWZambia,Stoc.12°SandWto¢.16°E. C.a.cordier:Rahm,1959—EDRCongoWoftheLualabaRiver,betweentheLowaandLuamariverstoclosetoLakeKivuintheEat¢.28°E(whereithybridizeswiththesubspeciesruwenzorii),andtothehighlandsWofLakeTanganyika,whereitextendstoc.6°S. C.a.cotton:Lydekker,1905—NEDRCongoEoftheCongo-L.ualabariversystem,fromtheNbankoftheCongoRiver(c.20°E)NtotheUeleRiver,EtoLakeAlbert,andStotheLindiRiver;formsahybridzonewiththesubspeciesruwenzoriiintheSofitsdistribution. C.a.palliatusPeters,1868—SEKenyaandNETanzania. C.a.prigogineiVerheyen,1959—EDRCongo(MtKabobo). C.a.ruwenzoriiThomas,1901—SEDRCongo,SEUganda,WRwanda,WBurundi,andNWTanzania;alongbothsidesoftheWRift,fromtheSemlikiValley,theRuwenzoriMts,andMtKahuzitotheshoresofLakeTanganyika. C. a. sharpei Thomas, 1902 — C & S Tanzania (as far W as Lake Rukwa), N Malawi, NW Zambia, and possibly occurring in N Mozambique. Descriptive notes. Head—body 55-66 cm (males) and 48-59 cm (females), tail 76 92 cm (males) and 63-76 cm (females); weight 7.6-12.6 kg (males) and 6.4-9.2 kg (females). The Angolan Colobus is generally black and characterized by long white epaulettes on shoulders and fluffy white circumfacial hair, the latter forming laterally elongated tufts on cheeks that resolve into a crescent shape. Tail may be tufted in males (not tufted in females), with varying extents of white (sometimes entirely white or with white restricted to the tuft). Two subspecies groups are recognized. The first is characterized by large epaulettes and a white pubic band (“Sclater’s Angolan Colobus ,” C. a. angolensis ; “Peters’s Angolan Colobus ,” C. a. palliatus; “Adolf Friedrich’s Angolan Colobus ,” C. a. ruwenzorii; and “Sharpe’s Angolan Colobus ,” C. a. sharper). The other subspecies group has thin epaulettes and no white band on the pubic region (“Cordier’s Angolan Colobus ,” C. a. cordieri; “Powell-Cotton’s Angolan Colobus ,” C. a. cottons; and “Prigogine’s Angolan Colobus ,” C. a. prigoginei). These two subspecies groups are linked by the unnamed Nkungwe Angolan Colobus that has well-developed epaulettes but lacks forehead and pubic bands. It is possible that some or all described subspecies may be distinct species. Sharpe’s Angolan Colobus is notable for its particularly long epaulettes, extending to its forearms, and bushy white side-whiskers. Prigogine’s Angolan Colobus has the smallest epaulettes and relatively reduced white side-whiskers. Habitat. A variety of forest types, from primary and secondary lowland rainforest and gallery forest to montane forest up to elevations of 3000 m and as high as 4000 m on Mount Ruwenzori. Although Angolan Colobus prefer the very highest levels of the forest canopy, including emergent trees, individuals will occasionally descend to the ground near streams to feed upon herbaceous vegetation. Food and Feeding. Diets of Angolan Colobus are mainly young leaves, along with shoots, seeds, unripe fruit, mature leaves, flowers, lichen, and termite clay. Water is obtained primarily from their food. Breeding. Female Angolan Colobus do not display a sexual swelling during the periovulatory period, and there is no perineal organ in the male. There is a definite birth season (May-December) in at least part of its distribution. A single young is born. The Natal coat is white, gradually changing through gray to black during the first six months. Activity patterns. The Angolan Colobus is diurnal and arboreal. At Nyungwe in Rwanda, its activity budget is 42% feeding, 32% resting, 20% moving, 5% engaging in social behavior, and 1% other activities. This population has unusually large group sizes (over 300 individuals), which may explain unusually high amounts of time spent feeding and moving and a relatively small amount of time devoted to resting, especially for a colobine. In a study in the Ituri forest of the DR Congo, groups of Angolan Colobus rested 43% of their time, fed 27%, moved 24%, engaged in social behavior 5%, and showed other behaviors 1%. Movements, Home range and Social organization. Daily movement has been reported at 983 m, with a home range of 371 ha. The Angolan Colobus generally lives in small family groups of 3—4 individuals, with a single male present. These family groups often form “troops” of up to 25-30 individuals, in which more than one male is present. One population in Rwanda (Nyungwe Forest) is exceptional in having a troop of more than 300 individuals, using a 2440ha home range. Little is known about the dispersal patterns of Angolan Colobus . Status and Conservation. CITES Appendix II. Classified as Least Concern on The IUCN Red List. The subspecies prigoginei is classified as Endangered, ruwenzorii as Vulnerable, cordieri as Near Threatened, and angolensis , cottoni, palliatus, and sharpei (under C. a. palliatus) as Least Concern. The undescribed Nkungwe taxon has not been assessed, but it is believed to be threatened. The Angolan Colobusis listed as Class B in the African Convention on the Conservation of Nature and Natural Resources. Although threatened in some parts ofits distribution (particularly in East Africa and the Albertine Rift), it remains widespread and relatively common in suitable habitat. Deforestation for timber and conversion of land for agricultural use are major threats, and isolated populations such as those in the Eastern Arc and coastal forests are especially vulnerable. It is also hunted for meat, especially in the Congo Basin. Populations in the eastern part ofits distribution are much more fragmented than those across the Congo Basin. The wide-ranging nominate form (Sclater’s Angolan Colobus ) is known to occur in Salonga National Park in DR Congo and Mweru Wantipa National Park in Zambia. Peters’s Angolan Colobus occurs in isolated forests in south-eastern Kenya, restricted to the southern coastal forests of the Kwale District; a comprehensive survey located 55 populations. Although locally threatened from habitat loss and hunting in some parts of its highly fragmented distribution, Peters’s Angolan Colobus remains widespread and relatively common. The total population is estimated at 3100-5000 individuals (560-900 groups), with the population in Shimba Hills Forest Reserve being the largest at ¢.2400 individuals (c.15 ind/km?). Sharpe’s Angolan Colobus is likewise discontinuously distributed in forest patches in the Southern Highlands and coastal and gallery forests of southern and eastern Tanzania. It occurs in Udzungwa Mountains National Park and Gendagenda South Forest Reserve. Cordier’s Angolan Colobus has declined as a result of increased hunting. It is not known whether it occurs in any protected areas. Powell-Cotton’s Angolan Colobus is locally threatened from habitat loss and hunting in some parts ofits distribution but remains widespread. It occurs in Kahuzi-Biéga National Park, Virunga National Park, and Tayna Gorilla Reserve in DR Congo. Prigogine’s Angolan Colobus is confined entirely to Mount Kabobo in eastern DR Congo, where it is at continuing risk of habitat loss from agricultural expansion and hunting. It is not known whether it occurs in any protected areas. Adolf Friedrich’s Angolan Colobus is restricted to a few sites in the Albertine Rift region, where its habitat is fragmented and at risk from continued degradation. Hunting is also a threat. It occurs in Ruwenzori National Park in Uganda, Nyungwe Forest National Park in Rwanda, Kaya Gonja Nature Reserve in Kenya, and Mahale Mountains National Park in Tanzania. Bibliography. Anderson, Cowlishaw & Rowcliffe (2007), Anderson, Rowcliffe & Cowlishaw (2007), Ansell (1959), Butynski & de Jong (2009), Fashing (2011), Fashing, Mulindahabi et al. (2007), Fimbel et al. (2001), Groves (1973, 2001, 2007b), Kanga & Heidi (1999/2000), Kingdon (1971, 1997), McDonald & Hamilton (2010), Moreno-Black & Bent (1982), Napier (1985), Nishida et al. (1981), Rahm (1970), Rodgers (1981), Verheyen (1959).